Response of sugar maple to calcium addition to northern hardwood forest

Watershed budget studies at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA, have demonstrated high calcium depletion of soil during the 20th century due, in part, to acid deposition. Over the past 25 years, tree growth (especially for sugar maple) has declined on the experimental watersheds at the HBEF. In October 1999, 0.85 Mg Ca/ha was added to Watershed 1 (W1) at the HBEF in the form of wollastonite (CaSiO3), a treatment that, by summer 2002, had raised the pH in the Oie horizon from 3.8 to 5.0 and, in the Oa horizon, from 3.9 to 4.2. We measured the response of sugar maple to the calcium fertilization treatment on W1. Foliar calcium concentration of canopy sugar maples in W1 increased markedly beginning the second year after treatment, and foliar manganese declined in years four and five. By 2005, the crown condition of sugar maple was much healthier in the treated watershed as compared with the untreated reference watershed (W6). Following high seed production in 2000 and 2002, the density of sugar maple seedlings increased significantly on W1 in comparison with W6 in 2001 and 2003. Survivorship of the 2003 cohort through July 2005 was much higher on W1 (36.6%) than W6 (10.2%). In 2003, sugar maple germinants on W1 were approximately 50% larger than those in reference plots, and foliar chlorophyll concentrations were significantly greater (0.27 g/m2 vs. 0.23 g/m2 leaf area). Foliage and fine-root calcium concentrations were roughly twice as high, and manganese concentrations twice as low in the treated than the reference seedlings in 2003 and 2004. Mycorrhizal colonization of seedlings was also much greater in the treated (22.4% of root length) than the reference sites (4.4%). A similar, though less dramatic, difference was observed for mycorrhizal colonization of mature sugar maples (56% vs. 35%). These results reinforce and extend other regional observations that sugar maple decline in the northeastern United States and southern Canada is caused in part by anthropogenic effects on soil calcium status, but the causal interactions among inorganic nutrition, physiological stress, mycorrhizal colonization, and seedling growth and health remain to be established.     

Soil responses to management, increased precipitation, and added nitrogen in ponderosa pine forests.

Forest management, climatic change, and atmospheric N deposition can affect soil biogeochemistry, but their combined effects are not well understood. We examined the effects of water and N amendments and forest thinning and burning on soil N pools and fluxes in ponderosa pine forests near Flagstaff, Arizona (USA). Using a 15N-depleted fertilizer, we also documented the distribution of added N into soil N pools. Because thinning and burning can increase soil water content and N availability, we hypothesized that these changes would alleviate water and N limitation of soil processes, causing smaller responses to added N and water in the restored stand. We found little support for this hypothesis. Responses of fine root biomass, potential net N mineralization, and the soil microbial N to water and N amendments were mostly unaffected by stand management. Most of the soil processes we examined were limited by N and water, and the increased N and soil water availability caused by forest restoration was insufficient to alleviate these limitations. For example, N addition caused a larger increase in potential net nitrification in the restored stand, and at a given level of soil N availability, N addition had a larger effect on soil microbial N in the restored stand. Possibly, forest restoration increased the availability of some other limiting resource, amplifying responses to added N and water. Tracer N recoveries in roots and in the forest floor were lower in the restored stand. Natural abundance delta15N of labile soil N pools were higher in the restored stand, consistent with a more open N cycle. We conclude that thinning and burning open up the N cycle, at least in the short-term, and that these changes are amplified by enhanced precipitation and N additions. Our results suggest that thinning and burning in ponderosa pine forests will not increase their resistance to changes in soil N dynamics resulting from increased atmospheric N deposition or increased precipitation due to climatic change. Restoration plans should consider the potential impact on long-term forest productivity of greater N losses from a more open N cycle, especially during the period immediately after thinning and burning.     

Demography of northern flying squirrels informs ecosystem management of western interior forests.

We studied northern flying squirrel (Glaucomys sabrinus) demography in the eastern Washington Cascade Range to test hypotheses about regional and local abundance patterns and to inform managers of the possible effects of fire and fuels management on flying squirrels. We quantified habitat characteristics and squirrel density, population trends, and demography in three typical forest cover types over a four-year period. We had 2034 captures of flying squirrels over 41 000 trap nights from 1997 through 2000 and marked 879 squirrels for mark-recapture population analysis. Ponderosa pine (Pinus ponderosa) forest appeared to be poorer habitat for flying squirrels than young or mature mixed-conifer forest. About 35% fewer individuals were captured in open pine forest than in dry mixed-conifer Douglas-fir (Pseudotsuga menziesii) and grand fir (Abies grandis) forests. Home ranges were 85% larger in pine forest (4.6 ha) than in mixed-conifer forests (2.5 ha). Similarly, population density (Huggins estimator) in ponderosa pine forest was half (1.1 squirrels/ha) that of mixed-conifer forest (2.2 squirrels/ha). Tree canopy cover was the single best correlate of squirrel density (r = 0.77), with an apparent threshold of 55% canopy cover separating stands with low- from high-density populations. Pradel estimates of annual recruitment were lower in open pine (0.28) than in young (0.35) and mature (0.37) forest. High recruitment was most strongly associated with high understory plant species richness and truffle biomass. Annual survival rates ranged from 45% to 59% and did not vary among cover types. Survival was most strongly associated with understory species richness and forage lichen biomass. Maximum snow depth had a strong negative effect on survival. Rate of per capita increase showed a density-dependent response. Thinning and prescribed burning in ponderosa pine and dry mixed conifer forests to restore stable fire regimes and forest structure might reduce flying squirrel densities at stand levels by reducing forest canopy, woody debris, and the diversity or biomass of understory plants, truffles, and lichens. Those impacts might be ameliorated by patchy harvesting and the retention of large trees, woody debris, and mistletoe brooms. Negative stand-level impacts would be traded for increased resistance and resilience of dry-forest landscapes to now-common, large-scale stand replacement fires.     

Influences of a calcium gradient on soil inorganic nitrogen in the Adirondack Mountains, New York

Studies of the long-term impacts of acidic deposition in Europe and North America have prompted growing interest in understanding the dynamics linking the nitrogen (N) and calcium (Ca) cycles in forested watersheds. While it has been shown that increasing concentrations of nitrate (NO3-) through atmospheric deposition or through nitrification can increase Ca loss, the reciprocal effects of Ca on N transformation processes have received less attention. We studied the influence of soil Ca availability on extractable inorganic N (NO3- + NH4+) across a Ca gradient in the Adirondack Mountains, New York, USA. Our results did not show the direct Ca-N interaction that we had expected, but instead showed that exchangeable Ca coupled with soil moisture, soil organic matter, and ambient temperature accounted for 61% of the variability in extractable inorganic N across 11 sites over two growing seasons. Soil Ca concentrations were, however, positively related to sugar maple (Acer saccharum) and American basswood (Tilia americana) basal areas and negatively related to American beech (Fagus grandifolia) basal area. Based on litter chemistry differences among these tree species and reported potential N mineralization values, we suggest that the influence of Ca on soil inorganic N is through a multistep pathway: reciprocal interactions between soil Ca concentrations and species composition, which in turn affect the quality of litter available for N mineralization. If chronic soil Ca depletion continues, as reported in some forested ecosystems, potential shifts in biotic communities could result in considerable alterations of N cycling processes.     

间伐对生态公益林冠层结构及土壤养分的影响

为定量分析上海城郊生态公益林间伐后林分冠层结构变化及对林下环境的影响,2009年7月采用TRAC植物冠层分析仪研究了香樟（Cinnamomum camphora）、杜英（Elaeocarpus sylvestris）和栾树（Koelreuteria paniculata）3种生态公益林间伐2年后林分冠层结构参数,设置临时样方调查各林分林下植物分布及生物量,测定了土壤N、P、K和有机质的变化。结果表明：间伐后3种林分的LAI、丛生指数和平均接触次数均降低,而间隙分数增大。间伐增加了林分间隙和LAI的空间异质性,间伐后3个树种LAI变异系数增加了19%～45%。轻度间伐后香樟、杜英和栾树林下PPFD（Photosynthetic Photon Flux Density）分别是未间伐林分的1.6、7和2.1倍,重度间伐香樟林内PPFD增大了8.9倍。间伐后林下植物种类、总盖度和干生物量随林下PPFD增加呈增大趋势。除全P和全K含量没有变化外,林地土壤全N、速效K、速效P和有机质含量均比未间伐林地有不同程度降低,但轻度间伐杜英林地速效养分消耗显著高于香樟和栾树林分。因此,研究区生态公益林结构调控需根据树种特性和林下环境变化,确定适宜的发展模式,提高公益林的综合效益。     

The spatial pattern of nitrogen cycling in the Adirondack Park, New York

Maps of canopy nitrogen obtained through analysis of high-resolution, hyperspectral, remotely sensed images now offer a powerful means to make landscape-scale to regional-scale estimates of forest N cycling and net primary production (NPP). Moreover, recent research has suggested that the spatial variability within maps of canopy N may be driven by environmental gradients in such features as historic forest disturbance, temperature, species composition, moisture, geology, and atmospheric N deposition. Using the wide variation in these six features found within the diverse forest ecosystems of the 2.5 million ha Adirondack Park, New York, USA, we examined linkages among environmental gradients and three measures of N cycling collected during the 2003 growing season: (1) field survey of canopy N, (2) field survey of soil C:N, and (3) canopy N measured through analysis of two 185 x 7.5 km Hyperion hyperspectral images. These three measures of N cycling strongly related to forest type but related poorly to all other environmental gradients. Further analysis revealed that the spatial pattern in N cycling appears to have distinct inter- and intraspecific components of variability. The interspecific component, or the proportional contribution of species functional traits to canopy biomass, explained 93% of spatial variability within the field canopy N survey and 37% of variability within the soil C:N survey. Residual analysis revealed that N deposition accounted for an additional 2% of variability in soil C:N, and N deposition and historical forest disturbance accounted for an additional 2.8% of variability in canopy N. Given our finding that 95.8% of the variability in the field canopy N survey could be attributed to variation in the physical environment, our research suggests that remotely sensed maps of canopy N may be useful not only to assess the spatial variability in N cycling and NPP, but also to unravel the relative importance of their multiple controlling factors.     

土壤酸化对油松生长的影响

褐色森林土酸化后,pH值下降,同时淋溶出大量Al离子和Mn离子。播种在酸化土壤中的油松种子萌发率稍有提高。生长在酸化土壤的油松苗的叶绿素含量,蒸腾强度,光合速率,相对生长率和干重均下降,原因是Al离子和Mn离子对油松的有毒害作用,并且阻止油松对钙离子和镁离子的吸收。表3参12     

东北过伐林区不同林分类型土壤肥力质量评价研究

保持和提高森林土壤肥力质量是实现森林健康的基础。研究不同林型下土壤肥力质量对森林可持续经营具有重要意义。以东北过伐林区金仓林场中的落叶松Lartx gmelinii人工纯林、云杉Picea asperatax紫椴Tilia amurensis天然混交林、紫椴×白桦Betula platyphylla天然混交林、五角槭Acer monox白桦×落叶松天然混交林、五角槭×白桦×紫椴天然混交林、白桦×落叶松天然混交林和五角械×紫椴×青杨Populus pseudo-simonii天然混交林为研究对象,分析和比较了不同林分下的土壤物理和化学性质,并采用主成分分析与聚类分析相结合的方法评价了其土壤肥力质量。结果表明：①林分类型对土壤理化性质影响显著;随土壤深度增加,土壤密度和土壤pH值增大,而土壤含水量、物理性粘粒含量和养分质量分数减少,但其在不同林分下的变化程度不同;②土壤因子之间关系密切,物理性粘粒含量与含水量、全氮、速效钾质量分数呈极显著正相关,而与土壤密度和pH值呈显著负相关;土壤养分质量分数之间具有不同程度的显著正相关关系;③采用主成分分析法对不同林分下0～60cm的土壤肥力质量进行了评价,土壤肥力质量表现为天然混交林高于落叶松人工纯林;采伐降低了土壤的肥力质量;在落叶松人工纯林中,随着林龄的增加,土壤的肥力质量下降。通过对土壤肥力质量综合指标值的聚类分析,将研究区土壤肥力质量分为优、良、中、差4个等级,肥力质量属于中等以上（优、良、中等）的林地所占研究样地的比例为66．7％,研究区森林土壤肥力质量属于中等水平。建议在经营过伐林时,注意调整林分结构和树种组成,控制采伐强度,同时进行长期定位观测和比较,以改善林分整体的肥力状况。本研究结果为该地区林业可持续经营提供直接依据,也为东北地区森林土壤肥力质量评价提供参考。     

No evidence that chronic nitrogen additions increase photosynthesis in mature sugar maple forests

Atmospheric nitrogen (N) deposition can increase forest growth. Because N deposition commonly increases foliar N concentrations, it is thought that this increase in forest growth is a consequence of enhanced leaf-level photosynthesis. However, tests of this mechanism have been infrequent, and increases in photosynthesis have not been consistently observed in mature forests subject to chronic N deposition. In four mature northern hardwood forests in the north-central United States, chronic N additions (30 kg N ha(-1) yr(-1) as NaNO3 for 14 years) have increased aboveground growth but have not affected canopy leaf biomass or leaf area index. In order to understand the mechanism behind the increases in growth, we hypothesized that the NO3(-) additions increased foliar N concentrations and leaf-level photosynthesis in the dominant species in these forests (sugar maple, Acer saccharum). The NO3(-) additions significantly increased foliar N. However, there was no significant difference between the ambient and +NO3(-) treatments in two seasons (2006-2007) of instantaneous measurements of photosynthesis from either canopy towers or excised branches. In measurements on excised branches, photosynthetic nitrogen use efficiency (micromol CO2 s(-1) g(-1) N) was significantly decreased (-13%) by NO3(-) additions. Furthermore, we found no consistent NO3(-) effect across all sites in either current foliage or leaf litter collected annually throughout the study (1993-2007) and analyzed for delta 13C and delta 18O, isotopes that can be used together to integrate changes in photosynthesis over time. We observed a small but significant NO3(-) effect on the average area and mass of individual leaves from the excised branches, but these differences varied by site and were countered by changes in leaf number. These photosynthesis and leaf area data together suggest that NO3(-) additions have not stimulated photosynthesis. There is no evidence that nutrient deficiencies have developed at these sites, so unlike other studies of photosynthesis in N-saturated forests, we cannot attribute the lack of a stimulation of photosynthesis to nutrient limitations. Rather than increases in C assimilation, the observed increases in aboveground growth at our study sites are more likely due to shifts in C allocation.     

不同林龄马尾松（Pinus massoniana）人工林碳氮磷分配格局及化学计量特征

为了解长江上游低山丘陵区马尾松（Pinus massoniana）人工林生态系统的C、N、P分配格局及化学计量特征,本文采用时空互代的方法,在宜宾高县来复林区选取三种不同林龄（5年生幼龄林、14年生中龄林、39年生成熟林）,但立地条件相近、样地情况基本一致的马尾松（Pinus massoniana）人工林作为研究对象,对马尾松针叶、凋落物及土壤中的C、N、P含量及 w（C）？w（N）？w（P）化学计量特征进行测定和分析。结果表明,（1）C、N、P 含量均表现为针叶〉凋落物〉土壤,且在三个库之间差异显著;（2）林龄对针叶、凋落物、土壤的 C、N、P 及 w（C）？w（N）、w（C）？w（P）计量比均有显著影响。（3）土壤 C、N、P含量在成熟林中最高;针叶和凋落物的C含量在成熟林中最低,N、P含量则在中龄林中最高。（4）随林龄增加马尾松对N、P的利用效率降低,针叶、凋落物及土壤的w（C）？w（N）与 w（C）？w（P）均表现为下降。（5）马尾松针叶w（N）？w（P）比值在14.37~15.53之间,说明该地区马尾松人工林受N和P的共同限制,但林龄对N、P养分限制的影响不显著。为提高该区马尾松人工林的生产力,建议在人工林的抚育管理中要适当增加N肥和P肥,同时也可在马尾松人工林引入豆科固氮植物以提高地力。该研究将马尾松针叶、凋落物及土壤结合起来探究随林龄增长C、N、P养分元素的分配格局及化学计量特征的变化,有助于全面、系统地揭示马尾松人工林生态系统的养分循环,对指导马尾松人工林生产,调节和改善林木生长环境,提高系统的养分利用效率及林地生产力具有重要意义。     

The role of canopy structural complexity in wood net primary production of a maturing northern deciduous forest

The even-aged northern hardwood forests of the Upper Great Lakes Region are undergoing an ecological transition during which structural and biotic complexity is increasing. Early-successional aspen (Populus spp.) and birch (Betula papyrifera) are senescing at an accelerating rate and are being replaced by middle-successional species including northern red oak (Quercus rubra), red maple (Acer rubrum), and white pine (Pinus strobus). Canopy structural complexity may increase due to forest age, canopy disturbances, and changing species diversity. More structurally complex canopies may enhance carbon (C) sequestration in old forests. We hypothesize that these biotic and structural alterations will result in increased structural complexity of the maturing canopy with implications for forest C uptake. At the University of Michigan Biological Station (UMBS), we combined a decade of observations of net primary productivity (NPP), leaf area index (LAI), site index, canopy tree-species diversity, and stand age with canopy structure measurements made with portable canopy lidar (PCL) in 30 forested plots. We then evaluated the relative impact of stand characteristics on productivity through succession using data collected over a nine-year period. We found that effects of canopy structural complexity on wood NPP (NPPw) were similar in magnitude to the effects of total leaf area and site quality. Furthermore, our results suggest that the effect of stand age on NPPw is mediated primarily through its effect on canopy structural complexity. Stand-level diversity of canopy-tree species was not significantly related to either canopy structure or NPPw. We conclude that increasing canopy structural complexity provides a mechanism for the potential maintenance of productivity in aging forests.     

川西高山森林生态系统林下生物量及其随林窗的变化特征

作为森林生态系统的重要组成部分,林下植被及其残体的分布受到林冠层的影响,但迄今有关林窗对林下植被和残体生物量的影响尚无研究报道.于2013 年8 月2 日至20 日,以海拔3 600 m 的川西岷江冷杉原始林林下植被为研究对象,根据区域内的坡向和林分组成等因素设置3 个100 m×100 m 的典型样地,调查其生物量及其随林窗的变化特征.在每个样地内选择3 个大林窗,在林窗、林缘和林下分别设置3 个20 m×20 m 的样方,调查粗木质残体长度或高度、大小头直径、枯立木记录胸径、腐烂等级等;在林窗、林缘和林下分别设置3 个5 m×5 m 的样方,采用“收获法”收集样方内直径在2.5-10 cm之间的细木质残体和灌木生物量;在林窗、林缘和林下分别设置3 个1 m×1 m 的样方来调查凋落物储量和草本生物量;在1m×1 m 的样方内随机选择1 个20 cm×20 cm 的小样方来调查地被植物生物量.结果表明,（1）川西高山森林生态系统总生物量为72.75 t·hm^-2,其中林下生物量为67.92 t·hm^-2,占生态系统生物量的95.17%.活体植被以灌木为主,其生物量为9.81t·hm^-2;残体部分以粗木质残体为主,其储量为53.00 t·hm^-2;（2）林窗对灌木、草本、地被植物的影响各不相同,且不同物种的灌木生物量表现出不同的分布规律;草本生物量表现出明显的“边缘效应”,在林缘显著高于林下;林窗和林缘的地被植物生物量相对较低;（3）粗木质残体储量从林下到林窗呈现减小的趋势,但总体储量仍然较大,林窗和林缘的细木质残体储量高于林下.这些结果为认识高山森林生态系统林下生物量及其格局,以及林窗在森林生态系统的重要作用提供了基础理论依据.     

Element mobility and partitioning along a soil acidity gradient in central Ontario forests, Canada

The potential environmental risk posed by metals in forest soils is typically evaluated by modeling metal mobility using soil-solution partitioning coefficients (K(d)), although such information is generally restricted to a few well-studied metals. Soil-solution partitioning coefficients were determined for 17 mineral elements (Al, As, Be, Ba, Ca, Cr, Cu, Fe, Ga, K, Li, Mg, Rb, Sr, Tl, U and V) in A-horizon (0-5 cm) soil at 46 forested sites that border the Precambrian Shield in central Ontario, where soil pH(aq) varied from 3.9 to 8.1. Sites were dominated by mature sugar maple (Acer saccharum Marsh.), white birch (Betula papyrifera Marsh.), balsam fir (Abies balsamea (L.) Mill.) or white pine (Pinus strobus L.). Log K(d) values for all elements could be predicted by empirical linear regression with soil pH (r (2) = 0.17-0.77) independent of forest type, although this relationship was greatly affected by positive relationships between acid-extractable metal concentration and pH(aq) for 13 of the 17 elements. Elements that exhibited strong or moderate (r (2)> 0.29; p < 0.001) relationships with soil pH(aq) in soil water extracts include Al, Ba, Fe, Ga, K, Li, Rb, Tl, V (negative) and Ca (positive). Elemental partitioning in mineral soil was independent of forest type; tree species differed in their response to chemical differences in mineral soil. For example, Rb, Ba, and Sr concentrations in foliage of sugar maple and white birch significantly increased with increasing soil acidity, whereas Rb, Ba, and Sr concentrations in balsam fir and white pine foliage exhibited no response to soil pH(aq). While K(d) values can provide useful information on the potential mobility and bioavailability of mineral elements in forest soils, care must be used when interpreting the relative contribution of solid and aqueous phases to this relationship and the differing responses of vegetation in elemental cycling in forests must also be considered.     

贵州麻江马尾松天然中龄林施肥研究

采用正交设计法对贵州麻江马尾松天然中龄林进行施肥试验，结果表明，施肥对林木生长具有良好的效果．至第７年，单株材积平均为对照的１３２．３％；所施元素中Ｎ、Ｐ、Ｋ对单株材积均有很大影响，且都达到０．１以上显著水平；而施Ｐ的自第４年至第７年均有良好的反应，其中以Ｐ２的效果最好；施肥的最优化组合为Ｎ２Ｐ２Ｋ２．     

Long-term CO2 enrichment of a forest ecosystem: implications for forest regeneration and succession.

The composition and successional status of a forest affect carbon storage and net ecosystem productivity, yet it remains unclear whether elevated atmospheric carbon dioxide (CO2) will impact rates and trajectories of forest succession. We examined how CO2 enrichment (+200 microL CO2/L air differential) affects forest succession through growth and survivorship of tree seedlings, as part of the Duke Forest free-air CO2 enrichment (FACE) experiment in North Carolina, USA. We planted 2352 seedlings of 14 species in the low light forest understory and determined effects of elevated CO2 on individual plant growth, survival, and total sample biomass accumulation, an integrator of plant growth and survivorship over time, for six years. We used a hierarchical Bayes framework to accommodate the uncertainty associated with the availability of light and the variability in growth among individual plants. We found that most species did not exhibit strong responses to CO2. Ulmus alata (+21%), Quercus alba (+9.5%), and nitrogen-fixing Robinia pseudoacacia (+230%) exhibited greater mean annual relative growth rates under elevated CO2 than under ambient conditions. The effects of CO2 were small relative to variability within populations; however, some species grew better under low light conditions when exposed to elevated CO2 than they did under ambient conditions. These species include shade-intolerant Liriodendron tulipifera and Liquidambar styraciflua, intermediate-tolerant Quercus velutina, and shade-tolerant Acer barbatum, A. rubrum, Prunus serotina, Ulmus alata, and Cercis canadensis. Contrary to our expectation, shade-intolerant trees did not survive better with CO2 enrichment, and population-scale responses to CO2 were influenced by survival probabilities in low light. CO2 enrichment did not increase rates of sample biomass accumulation for most species, but it did stimulate biomass growth of shade-tolerant taxa, particularly Acer barbatum and Ulmus alata. Our data suggest a small CO2 fertilization effect on tree productivity, and the possibility of reduced carbon accumulation rates relative to today's forests due to changes in species composition.     

Seeing the forest for the heterogeneous trees: stand-scale resource distributions emerge from tree-scale structure

Forest ecosystem processes depend on local interactions that are modified by the spatial pattern of trees and resources. Effects of resource supplies on processes such as regeneration are increasingly well understood, yet we have few tools to compare resource heterogeneity among forests that differ in structural complexity. We used a neighborhood approach to examine understory light and nutrient availability in a well-replicated and large-scale variable-retention harvesting experiment in a red pine forest in Minnesota, USA. The experiment included an unharvested control and three harvesting treatments with similar tree abundance but different patterns of retention (evenly dispersed as well as aggregated retention achieved by cutting 0.1- or 0.3-ha gaps). We measured light and soil nutrients across all treatments and mapped trees around each sample point to develop an index of neighborhood effects (NI). Field data and simulation modeling were used to test hypotheses that the mean and heterogeneity of resource availability would increase with patchiness because of greater variation in competitive environments. Our treatments dramatically altered the types and abundances of competitive neighborhoods (NI) in each stand and resulted in significantly nonlinear relationships of light, nitrogen and phosphorus availability to NI. Hence, the distribution of neighborhoods in each treatment had a significant impact on resource availability and heterogeneity. In dense control stands, neighborhood variation had little impact on resource availability, whereas in more open stands (retention treatments), it had large effects on light and modest effects on soil nutrients. Our results demonstrate that tree spatial pattern can affect resource availability and heterogeneity in explainable and predictable ways, and that neighborhood models provide a useful tool for scaling heterogeneity from the individual tree to the stand. These insights are needed to anticipate the outcomes of silvicultural manipulations and should become more holistically integrated into both basic ecological and management science.     

Shifts in southern Wisconsin forest canopy and understory richness, composition, and heterogeneity

We resurveyed the under- and overstory species composition of 94 upland forest stands in southern Wisconsin in 2002-2004 to assess shifts in canopy and understory richness, composition, and heterogeneity relative to the original surveys in 1949-1950. The canopy has shifted from mostly oaks (Quercus spp.) toward more mesic and shade-tolerant trees (primarily Acer spp.). Oak-dominated early-successional stands and those on coarse, nutrient-poor soils changed the most in canopy composition. Understories at most sites (80%) lost native species, with mean species density declining 25% at the 1-m2 scale and 23.1% at the 20-m2 scale. Woody species have increased 15% relative to herbaceous species in the understory despite declining in absolute abundance. Initial canopy composition, particularly the abundance of red oaks (Quercus rubra and Q. velutina), predicted understory changes better than the changes observed in the overstory. Overall rates of native species loss were greater in later-successional stands, a pattern driven by differential immigration rather than differential extirpation. However, understory species initially found in early-successional habitats declined the most, particularly remnant savanna taxa with narrow or thick leaves. These losses have yet to be offset by compensating increases in native shade-adapted species. Exotic species have proliferated in prevalence (from 13 to 76 stands) and relative abundance (from 1.2% to 8.4%), but these increases appear unrelated to the declines in native species richness and heterogeneity observed. Although canopy succession has clearly influenced shifts in understory composition and diversity, the magnitude of native species declines and failure to recruit more shade-adapted species suggest that other factors now act to limit the richness, heterogeneity, and composition of these communities.     

Nutrient limitation in soils exhibiting differing nitrogen availabilities: what lies beyond nitrogen saturation?

The nature of nutrient limitation in large areas of temperate forest may be changing due to human activities. As N availability in these forests increases, other nutrients could increasingly constrain productivity and other ecosystem processes. To determine the nature of nutrient limitation (N, P, and Ca) in forest soils exhibiting differing N availability, we conducted three field studies in the Fernow Experimental Forest, West Virginia, USA. The first used a ubiquitous herbaceous species, Viola rotundifolia, to compare indices of N availability to the activity of root-associated phosphomonoesterase (PME) activity at two spatial scales. The second study used fertilized, root in-growth cores to assess the extent of N, P, and Ca limitation. Finally, we measured the root-associated PME activity of V. rotundifolia growing in experimental plots that have received various combinations of nutrient additions and harvest treatments. For entire watersheds, stream water nitrate concentrations were positively related to PME activities (R2 = 0.986). For small plots, PME activities were positively associated with soil nitrate availability (R2 = 0.425), and to a lesser extent with the leaf N concentrations (R2 = 0.291). Root growth into microsites fertilized with P was greater than growth into microsites fertilized with either N or Ca, especially in watersheds with high N availability. Experimental additions of N increased the root-associated PME activity of V. rotundifolia, supporting the causality of the relationship between N availability and PME activity. Collectively, our results indicate that, as N availability increases, P becomes increasingly limiting at the sites examined. Understanding how nutrient limitations change during N saturation should improve ecosystem models and better inform our attempts to mitigate any undesired effects.     

Seedling growth responses to soil resources in the understory of a wet tropical forest

Plant growth responses to resources may be an important mechanism that influences species' distributions, coexistence, and community structure. Irradiance is considered the most important resource for seedling growth in the understory of wet tropical forests, but multiple soil nutrients and species have yet to be examined simultaneously with irradiance under field conditions. To identify potentially limiting resources, we modeled tree seedling growth as a function of irradiance and soil nutrients across five sites, spanning a soil fertility gradient in old-growth, wet tropical forests at La Selva Biological Station, Costa Rica. We measured an array of soil nutrients including total nitrogen (total N), inorganic N (nitrate [NO3-] and ammonium [NH4+]), phosphate (PO4-), and sum of base cations (SBC; potassium, magnesium, and calcium). Shade in the forest understory did not preclude seedling growth correlations with soil nutrients. Irradiance was a significant predictor of growth in 52% of the species, inorganic N in 54% (NO3- in 32%; NH4+ in 34%), total N in 47%, SBC in 39%, and PO4- in 29%. Overall, growth was correlated with both irradiance and soil nutrients in 45% of species and with soil nutrients only in an additional 48%; rarely was irradiance alone correlated with growth. Contrary to expectations, the magnitudes of growth effects, assessed as the maximum growth response to significant resources for each species, were similar for irradiance and most soil nutrients. Among species whose growth correlated with soil nutrients, the rank importance of nutrient effects was SBC, followed by N (total N, NO3-, and/or NH4+) and PO4-. Species' growth responsiveness (i.e., magnitudes of effect) to irradiance and soil nutrients was negatively correlated with species' shade tolerance (survival under 1% full sun). In this broad survey of species and resources, the nearly ubiquitous effects of soil nutrients on seedling growth challenge the idea that soil nutrients are less important than irradiance in the light-limited understory of wet tropical forests.     

Linkages between phenology, pollination, photosynthesis, and reproduction in deciduous forest understory plants

Light availability in the understory of deciduous forests changes drastically within the growing season due to the foliage dynamics of canopy trees. Because flowering phenology, photosynthetic characteristics, and fruiting success respond to such strong seasonality in light availability, we hypothesized that understory plants in such ecosystems should describe distinct phenological groups or syndromes where "syndrome" is defined only as a set of characteristics that co-occur. To identify these phenological syndromes, we studied the flowering phenology, fruit or seed set, and photosynthetic characteristics for 18 perennial understory herbaceous species that differed in reproductive strategy over eight years in a deciduous forest in northern Japan. Three phenological groups emerged from this study: (1) spring bloomers, flowering and fruiting before the completion of canopy closure; (2) early-summer bloomers, flowering during the progress of canopy closure and fruiting after canopy closure; and (3) late-summer bloomers, flowering and fruiting after canopy closure. The spring bloomers had high photosynthetic rates and high fruiting abilities, but the flowering time varied considerably among years due to yearly fluctuations of snowmelt date. Bumble bee-pollinated species of spring bloomers showed variable seed-set success, while fly-pollinated species showed relatively stable seed sets over the years. The early-summer bloomers showed low fruiting abilities irrespective of pollination success, reflecting severe resource limitation with decelerating light availability during fruit development. Although the late-summer bloomers showed low photosynthetic rates under low-light conditions, high fruit-set success was attained if pollination was sufficient. These results support our hypothesis that phenological syndromes may be found in deciduous forest understory plants. Given that reproductive success of bee-pollinated spring bloomers is highly susceptible to seasonal fluctuation, climate change may have its strongest impacts on this group.