Habitat niche specialization in an understory species in a warm temperate forest

Relationships between microhabitat variables; understory light conditions in summer and winter, altitude, slope inclination and topographic categories (valley, ridge, and slope) and the distribution of Aucuba japonica Thunb. (Cornaceae), a common understory shrub species in Japan were examined using non-contagious 66, 20 × 20 m² quadrats. The Morishita’s I _δ suggested that A. japonica distributions were strongly heterogeneous among the quadrats. Therefore positive spatial autocorrelation of A. japonica at a within-quadrat level (≤20 m) was obvious. Moran’s I statistics showed a significant positive spatial autocorrelation in A. japonica abundance within the distance shorter than 60 m. But the partial Mantel tests suggested that the mass effect from neighboring quadrats would little explain A. japonica abundance in a quadrat. The partial Mantel tests also clearly showed that A. japonica distributions were strongly structured by topography and understory light conditions. Using Monte Carlo randomization tests, we found that A. japonica was aggregately distributed in quadrats in valley which were covered by deciduous canopies. Better understory light conditions in winter together with valley edaphic conditions may increase the abundance of A. japonica there. It is concluded that habitat niche specialization is important in structuring distribution of A. japonica in this forest community.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Studies on the life history of an evergreen herb,Pyrola japonica,population on a forest floor in a warm temperate region

The life history and matter economy were studied on an evergreen herb,Pyrola japonica Klenze, population growing on the floor of a deciduous forest in a warm temperate region, central Japan. Seasonal changes in standing crop, bulk density and reserve substance of each organ and in leaf area were investigated over a year. Monthly and annual net production of the population were estimated based on the growth in dry weight and the seasonal dynamics of reserve substance. Seasonal peak value of the monthly net production was 16.7 g d.w.m⁻² in May. The annual net production was estimated to be 60.2 g d.w.m⁻². Two phases were recognized in the annual pattern of development and matter economy of this plant. Phase I, from April to June, was characterized by the development of aboveground vegetative organs and the consumption of reserve substance rrom old organs. Phase II, from July to March, was characterized by the development of reproductive and underground organs, and the accumulation of reserve substance. Phase II was further divided into two sub-phases according to the behavior of the reserve substance and the rate of net production. The production process of the population was compared with those of other evergreen herbs growing on the forest floor in warm temperate regions.     

暖温带落叶阔叶林动态变化的模拟研究

用森林动态林窗模型 FORET1模拟了暖温带落叶阔叶林的长期变化特征。模型参数取自暖温带地区长期森林研究和经营的历史数据 ,对过去数据中缺少的参数进行了实地测定 ,并用观测的数据对模型作了检验。结果表明模型能较好地模拟暖温带落叶阔叶林的长期动态变化特征。通过模拟可以看出 ,森林的净初级生产力没有明显变化规律且极度不稳定 ,峰值出现在30 a左右 ,相似于世界上其它地区森林动态格局变化 ,生物量格局呈循环状态变化 ,循环周期大致在 110 a左右。     

Studies on the life history of the population of an evergreen herb,Pyrola japonica,on the forest floor in a warm temperate region

Photosynthetic and respiratory activities and gross production in relation to temperature conditions were investigated in the population of an evergreen herb,Pyrola japonica, growing on the floor of a deciduous forest in the warm temperate region of central Japan. Analysis of the temperature-photosynthesis relationship ofP. japonica leaves during the growing season indicated distinct seasonal changes in the temperature optimum for photosynthesis. This population was found to be acclimatable to ambient air temperatures exceeding 15C, but this acclimation became less pronounced under thermal conditions below 15 C. This plant possessed narrow photosynthetic optima in the warm season but wide optima in the cold season. The shape of the temperature-respiration curve did not vary significantly with the months except for April. The Q₁₀ for respiration between 10 C and 20 C was calculated to be 1.93–2.65. Annual dry matter loss associated with respiration was estimated to amount to 159.1 g d.w.m⁻² based on the measurements of the seasonal changes in the respiratory activity of each organ. Gross production of this population was estimated to be 219.3 g d.w.m⁻² year⁻¹ as the sum total of the net production (60.2 g d.w.m⁻²year⁻¹) and the respiration. Monthly gross production was high in the early growing season, and low and stable in winter.     

Biomass and carbon storage of the North American deciduous forest

Field measures of tree and shrub dimensions were used with established biomass equations in a stratified, two-stage cluster sampling design to estimate above-ground ovendry woody biomass and carbon storage of the eastern deciduous forest of North America. Biomass averaged 8.1 ± 1.4 (95% C.I.) kg/m² and totaled 18.1 ± 3.1 (95% C.I.) gigatons. Carbon storage averaged 3.6 ± 0.6 (95% C.I.) kg/m² and totaled 8.1 ± 1.4 (95% C.I.) gigatons. These values are lower than previous estimates commonly used in the analysis of the global carbon budget which range from 17.1 to 23.1 kg/m² for biomass and 7.7 to 10.4 kg/m² for carbon storage. These new estimates for the deciduous forest, together with earlier work in the boreal forest begin to reveal a pattern of overestimation of global carbon storage by vegetation in analyses of the global carbon budget. We discuss reasons for the differences between the new and earlier estimates, as well as implications for our understanding of the global carbon cycle.     

Comparative study of leaf carbon gain in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool-temperate deciduous forest

Seasonal courses of leaf CO₂ gas exchange in a growing season were examined in saplings ofThujopsis dolabrata var.hondai andQuercus mongolica var.grosseserrata in a cool temperate deciduous forest. Between the two tree species there were no large differences in the light compensation point of leaf photosynthesis, except for the season of new leaf expansion. However, light-saturated rates of net photosynthesis were obviously high inT. dolabrata var.hondai. EvergreenT. dolabrata var.hondai saplings had large photosynthetic production in two seasons, before the emergence of new foliage and after foliage fall of the overstory deciduous trees, because of the significantly high solar radiant energy penetrating under the forest canopy during the seasons. Saplings of deciduousQ. mongolica var.grosseserrata were heavily shaded throughout the growing season by foliage of the overstory trees, which resulted in a low daily surplus production. The annual surplus production of leaves in the growing season was estimated to be 2300 mmol CO₂ m⁻² inT. dolabrata var.hondai and −100 mmol CO₂ m⁻², slightly negative, inQ. mongolica var.grosseserrata. These results supported the high survivability ofT. dolabrata var.hondai saplings and the high mortality ofQ. mongolica var.grosseserrata in the deciduous forest.     

Leaf carbon gain simulation for tree seedlings acclimatized to microsite light regimes of a temperate pine forest

Leaf carbon gain simulation was performed forQuercus serrata seedlings with previously reported 6 day photosynthetic photon flux density (PPFD) histograms from 20 understorey microsites of a pine forest (Washitani & Tang 1991). This simulation was performed with or without an assumption of the acclimatization of photosynthetic capacity (P_max) to microsite light availability. A constant ratio of respiration rate to P_max, within, the range of 0.07–0.1, was assumed as a constraint. In relatively well illuminated microsites with a diffuse site factor above 0.1, predicted optimal P_max was about 5 μmol m⁻² s⁻¹, with the predicted mean daily net carbon gain being about 50 mmol m⁻² day⁻¹. Each of the predicted optimal P_max and the simulated mean daily net carbon gains with a constant P_max (5 μmol m⁻² s⁻¹) or the predicted optimal P_max was linearly related to the microsite light availability index, diffuse site factor. Simulated net carbon gain was negative at diffuse site factors below 0.04, if the constant of P_max was assumed. The predicted linear relationship between net carbon gain and diffuse site factor could provide an ecophysiological basis for the observed linear dependency of the relative growth rate of biomass ofQ. serrata seedlings on the microsite diffuse site factor (Washitani & Tang 1991).     

Resource allocation to sexual and vegetative reproduction in a forest herbSyneilesis palmata (Compositae)

The balance between sexual and vegetative reproduction inSyneilesis palmata was examined in relation to environmental conditions and the amount of reproductive resources, which is defined here as the total quantity of dry matter invested in both modes of reproduction. The allocation balance was measured for individual plants of two populations, with different densities in an open habitat, for 2 years, and those of two other populations, under different light intensities in a plantation forest (forest floor and edge), for 3 years. Relative allocation to sexual reproduction decreased with increasing reproductive resources in all populations except for the forest edge, which showed a constant allocation balance. The high density population showed lower relative allocation to sexual reproduction than the low density population, irrespective of the amount of reproductive resources. However the between-year comparison of the high density population suggested that under extremely high density, plants with a small amount of reproductive resources enhanced sexual reproduction, while plants with a large amount of reproductive resources reproduced vegetatively. On the forest floor, plants with small amounts of reproductive resources had higher relative allocation to sexual reproduction than the forest edge population, while plants with large amounts of reproductive resources had a somewhat lower one. The adaptive significance of such allocation patterns are discussed, based on qualitative data on the characteristics of both types of offspring.     

Post‐clearcut dynamics of carbon,water and energy exchanges in a midlatitude temperate,deciduous broadleaf forest environment

Clearcutting and other forest disturbances perturb carbon, water, and energy balances in significant ways, with corresponding influences on Earth's climate system through biogeochemical and biogeophysical effects. Observations are needed to quantify the precise changes in these balances as they vary across diverse disturbances of different types, severities, and in various climate and ecosystem type settings. This study combines eddy covariance and micrometeorological measurements of surface‐atmosphere exchanges with vegetation inventories and chamber‐based estimates of soil respiration to quantify how carbon, water, and energy fluxes changed during the first 3 years following forest clearing in a temperate forest environment of the northeastern US. We observed rapid recovery with sustained increases in gross ecosystem productivity (GEP) over the first three growing seasons post‐clearing, coincident with large and relatively stable net emission of CO₂ because of overwhelmingly large ecosystem respiration. The rise in GEP was attributed to vegetation changes not environmental conditions (e.g., weather), but attribution to the expansion of leaf area vs. changes in vegetation composition remains unclear. Soil respiration was estimated to contribute 44% of total ecosystem respiration during summer months and coarse woody debris accounted for another 18%. Evapotranspiration also recovered rapidly and continued to rise across years with a corresponding decrease in sensible heat flux. Gross short‐wave and long‐wave radiative fluxes were stable across years except for strong wintertime dependence on snow covered conditions and corresponding variation in albedo. Overall, these findings underscore the highly dynamic nature of carbon and water exchanges and vegetation composition during the regrowth following a severe forest disturbance, and sheds light on both the magnitude of such changes and the underlying mechanisms with a unique example from a temperate, deciduous broadleaf forest.     

Early spring leaf out enhances growth and survival of saplings in a temperate deciduous forest

Saplings of many canopy tree species in winter deciduous forests receive the major portion of their light budget for their growing season prior to canopy closure in the spring. This period of high light may be critical for achieving a positive carbon (C) gain, thus contributing strongly to their growth and survival. This study of saplings of Aesculus glabra and Acer saccharum in Trelease Woods, Illinois, USA, tested this hypothesis experimentally by placing tents of shade cloth over saplings during their spring period of high light prior to canopy closure in three consecutive years. Leaf senescence began 16 days (year 0) and 60 days (year 1) earlier for shaded A. glabra saplings than control saplings. No change in senescence occurred for A. saccharum. The annual absolute growth in stem diameter of both species was negligible or negative for shaded saplings, but positive for control saplings. Only 7% of the shaded A. glabra saplings were alive after 2 years, while all control saplings survived for 3 years; only 20% of the shaded A. saccharum saplings survived for 3 years, while 73% of control saplings were alive after the same period. Early spring leaf out is a critical mechanism that allows the long-term persistence of saplings of these species in this winter deciduous forest. Studies and models of C gain, growth, and survival of saplings in deciduous forests may need to take into account their spring phenology because saplings of many species are actually “sun” individuals in the spring prior to their longer period in the summer shade.     

Effects of rainfall events on soil CO2 flux in a cool temperate deciduous broad-leaved forest

The effects of rainfall events on soil CO₂ fluxes were examined in a cool temperate Quercus/Betula forest in Japan. The soil CO₂ fluxes were measured using an open-flow gas exchange system with an infrared gas analyzer in the snow-free season from August 1999 to November 2000. Soil CO₂ flux showed no significant diurnal trend on days without rain. In contrast, rainfall events caused a significant increase in soil CO₂ flux. To determine the effect of rainfall events and to evaluate more precisely the daily and annual soil carbon flux, we constructed a multiple polynomial regression model that included two variables, soil temperature and soil water content, using the soil CO₂ flux data recorded on sunny days. Daily soil carbon fluxes on sunny days calculated by the model were almost the same as those determined by the field measurements. On the contrary, the fluxes measured on rainy days were significantly higher than those calculated daily from the soil carbon fluxes by the model. Annual soil carbon fluxes in 1999 and 2000 were estimated using models that both do and do not take rainfall effects into consideration. The result indicates that post-rainfall increases in soil CO₂ flux represent approximately 16–21% of the annual soil carbon flux in this cool temperate deciduous forest.     

Soil seed bank composition and diversity in a managed temperate deciduous forest

Little is known about the influence of forest management on the interaction between seed bank and aboveground vegetation. We surveyed seed banks and vegetation in 10 forest stands under similar abiotic conditions but submitted either to a coppice-with-standards treatment (n=5) or to a selective-cutting system (n=5). We analyzed species composition and diversity, community ecological profile, and distribution of taxa among different life forms, strategy, morphology and functional type categories. A total of 2085 seedlings (8296 seedsm^–2) germinated-corresponding to 28 species, among which Juncus effusus was the most abundant. Fifty-seven percent of the species were also recorded in the aboveground vegetation, the dominant species being Rubus fruticosus agg., but only 28% of the aboveground species were present in the seed bank. Our results suggest that (1) vernal geophytes and shade-tolerant perennials, which group most true forest species, are not incorporated in the seed bank, (2) parent plants of most seeds were present either in the stand in an earlier dynamic stage or apart from the stand and long-distance dispersed, (3) as expected, early-successional species are well represented in the seed bank, (4) forestry vehicles seem to be a major means of dispersion for stress-tolerant species normally found in forest lanes and wheel tracks. We conclude that seed banks contain species that have a potentially negative impact on the true forest flora and, thus, forest management should minimize soil disturbance and retain remnants of old-coppice woods to conserve disturbance-sensitive true forest species.     

Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy

Water-use strategies of Populus tremula and Tilia cordata, and the role of abscisic acid in these strategies, were analysed. P. tremula dominated in the overstorey and T. cordata in the lower layer of the tree canopy of the temperate deciduous forest canopy. Shoot water potential (), bulk-leaf abscisic acid concentration ([ABA]_leaf), abscisic acid concentration in xylem sap ([ABA]_xyl), and rate of stomatal closure following the supply of exogenous ABA (v) decreased acropetally through the whole tree canopy, and foliar water content per area (w), concentration of the leaf osmoticum (c), maximum leaf-specific hydraulic conductance of shoot (L), stomatal conductance (g_s), and the threshold dose per leaf area of the exogenous ABA (d_a) required to reduce stomatal conductance increased acropetally through the tree canopy (from the base of the foliage of T. cordata to the top of the foliage of P. tremula) in non-stressed trees. The threshold dose per leaf dry mass of the exogenous ABA (d_w) required to reduce stomatal conductance, was similar through the tree canopy. After a drought period (3 weeks), the , w, L, g_s, d_a and d_w had decreased, and c and v had increased in both species. Yet, the effect of the drought period was more pronounced on L, g_s, d_a, d_w and v in T. cordata, and on , w and c in P. tremula. It was concluded that the water use of the species of the lower canopy layer—T. cordata, is more conservative than that of the species of the overstorey, P. tremula. [ABA]_leaf had not been significantly changed in these trees, and [ABA]_xyl had increased during the drought period only in P. tremula. The relations between [ABA]_leaf, [ABA]_xyl and the stomatal conductance, the osmotic adjustment and the shoot hydraulic conductance are also discussed.     

Vegetation structure and diversity of an ancient temperate deciduous forest in SW Denmark

Bolderslev Skov (113 ha) is the largest contiguous ancient forest remnant in Denmark. The forest has been preserved since 1998 as a strict non-intervention forest reserve. We studied vegetation structure, floristic gradients, and diversity of the forest in 50 plots of 100 m² placed according to a restricted random sampling design. Dominant tree species were Fagus sylvatica, Fraxinus excelsior, Tilia cordata and Quercus robur, which in most parts of the forest form a mixed canopy. Most stands appeared to be of moderate age, 55-80 years old, and large old trees were rare. pH in association with light and thickness of the litter layer were the most important factors in explaining floristic gradients in the forest. Soil moisture (dry to mesic) was not strongly correlated with DCA axes for neither tree nor field layer, but had a significant effect on the distribution of a number of herb species. Forest structure was not important in explaining species distributions. Field layer species richness showed a positive relationship with the pH-gradient. At the scale of 1m² plots we also found a highly significant negative relationship between field layer species richness and the plot-wise Ellenberg indicator value for nitrogen availability. Structure of the tree layer had little effect on field layer species richness. The mixed composition, long continuity, and presence of a high proportion of the regional species pool assigns a high conservation value to Bolderslev Skov and makes it an important site for future studies of the dynamic properties, niche preferences, and inter-specific competition of temperate deciduous forest species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Seasonal food habits of a sika deer population in the warm temperate forest of the westernmost part of Honshu, Japan

The rumen contents of 63 sika deer (Cervus nippon Temminck) shot in the westernmost part of Honshu were analyzed by the point-frame method. The rumen contents were dominated by forbs and the majority of these were evergreen monocotyledonous plants, probably Ophiopogon spp. Evergreen browse leaves (woody leaves), such as Eurya japonica Thunb., Ilex crenata Thunb. and Trachelospermum asiaticum Nakai, were also important, constituting 20–40% of the rumen contents. Other categories accounted for only small proportions, although in summer, graminoids increased but twigs and bark decreased in importance and in fall and winter, fruits and seeds increased in importance. The diet of the deer was characterized by high-quality foods even in winter and by a stability or less-pronounced seasonality throughout the year than found in the northern deer. These results were compared with those from other populations, including a northern sika deer population in Japan.     

Shoot life span in relation to successional status in deciduous broad-leaved tree species in a temperate forest

In trees, leaf life span is closely related to successional status. Although leaves are attached to shoots, shoot life span has been insufficiently studied in the context of ecological systems. Interspecific variation in shoot survivorship was investigated over 27 months in 15 temperate hardwood tree species. Relationships between shoot architecture and shoot survival were also investigated. Shoot life span was shortest in early successional species, and longest in late successional species, in each of the families Betulaceae and Fagaceae. In Salicaceae, all of which were early successional species, shoot life span was longer in mountainous than in riparian species. Early successional or riparian species distributed longer shoots densely, even in proximal positions on mother shoots, resulting in mutual shading and consequent early and massive shoot shedding. By contrast, late successional or mountainous species concentrated shoots in distal positions, allowing shoots to receive equally favorable light, resulting in a longer life span. These results reveal close relationships between shoot life span and environmental resource availability or successional status and suggest a causal relationship between shoot shedding and shoot architecture.     

An initial intercomparison of micrometeorological and ecological inventory estimates of carbon exchange in a mid-latitude deciduous forest

The role of mid‐latitude forests in the sequestration of carbon (C) is of interest to an increasing number of scientists and policy‐makers alike. Net CO₂ exchange can be estimated on an annual basis, using eddy‐covariance techniques or from ecological inventories of C fluxes to and from a forest. Here we present an intercomparison of annual estimates of C exchange in a mixed hardwood forest in the Morgan‐Monroe State Forest, Indiana, USA for two years, 1998 and 1999. Based on eddy‐covariance measurements made at 1.8 times canopy height from a tower, C uptake by the forest was 237 and 287 g C m^?2 y^?1 for 1998 and 1999, respectively. For the same time period, biometric and ecophysiological measures and modelled estimates of all significant carbon fluxes within deciduous forests were made, including: change in living biomass, aboveground and belowground detritus production, foliage consumption, and forest floor and soil respiration. Using this ecological inventory method for these same two time periods, C uptake was estimated to be 271 and 377 g C m^?2 y^?1, which are 14.3% and 31.4% larger, respectively, than the tower‐based values. The relative change between this method's annual estimates is consistent with that of the eddy‐covariance based values. Our results indicate that the difference in annual C exchange rates was due to reduced heterotrophic soil respiration in 1999.     

Controls on the dynamics of dissolved organic carbon and nitrogen in a Central European deciduous forest

Despite growing attention concerning therole of dissolved organic matter (DOM) inelement cycling of forest ecosystems, thecontrols of concentrations and fluxes of bothdissolved organic carbon (DOC) and nitrogen(DON) under field conditions in forest soilsremain only poorly understood. The goal ofthis project is to measure the concentrations and fluxes of DON, NH₄ ⁺, NO₃ ^–and DOC in bulkprecipitation, throughfall, forest floorleachates and soil solutions of a deciduousstand in the Steigerwald region (northernBavaria, Germany). The DOC and DONconcentrations and fluxes were highest inleachates originating from the Oa layer of theforest floor (73 mg C L^–1, 2.3 mg NL^–1 and about 200–350 kg C, 8–10 kg Nha^–1 yr^–1). They were observed to behighly variable over time and decreased in themineral topsoil (17 mg C L^–1, 0.6 mg NL^–1 and about 50–90 kg C, 2.0 to 2.4 kg Nha^–1 yr^–1). The annual variability ofDOC and DON concentrations and subsequentialDOC/DON ratios was substantial in allsolutions. The DOC and DON concentrations inthroughfall were positively correlated withtemperature. The DOC and DON concentrationsdid not show seasonality in the forest floorand mineral soil. Concentrations were notrelated to litterfall dynamics but didcorrespond in part to the input of DOC and DONfrom throughfall. The throughfall contributionto the overall element fluxes was higher forDON than for DOC. Concentrations and fluxes ofDON were significantly correlated to DOC inthroughfall and the Oi layer. However, thecorrelation was weak in Oa leachates. Inaddition, seasonal and annual variation ofDOC/DON ratios indicated different mechanismsand release rates from the forest floor forboth components. The concentrations of DOC andDON in forest floor leachates were in mostcases dependent neither on the pH value orionic strength of the solution, nor on thewater flux or temperature changes. As aconsequence, the DOC and DON fluxes from theforest floor into the mineral soil werelargely dependent on the water flux if annualand biweekly time scales are considered.     

The spatial pattern and canopy-understory association of trees in a cool temperate,mixed forest in western Japan

The spatial distribution pattern of trees and the association between canopy and understory individuals were examined with reference to the distribution of tree crowns in a cool temperate, mixed forest in Ohdaigahara, western Japan. Line transect and contact sampling methods were used to examine the pattern over various spatial scales. These methods are useful to detect patterns over a large study area. The dominance ofChamaecyparis obtusa on steep slopes forming large patches suggested that the distribution of this species is a consequence of landslides. UnderstoryFagus crenata showed a clumped distribution, and the relative coverage of this species was larger in canopy gaps than under a closed canopy. Understory individuals ofAbies homolepis showed a positive association with canopy trees ofF. crenata but a negative association with conspecific canopy individuals. These patterns suggested thatF. crenata regenerates in canopy gaps and is replaced byA. homolepis. The dynamics of these two species are consistent with the process of gap dynamics. The effects of both small- and large-scale disturbance must be evaluated to understand the mechanisms of patch formation and the coexistence of forest tree species.