Nitrogen storage forms in nine boreal understorey plant species

Storage forms of N were studied in below-ground structures of nine boreal forest understorey plants. The ericaceous shrubs Vacciniumvitis-idaea and V.myrtillus, the fern Gymnocarpium dryopteris, the grass Deschampsia flexuosa, and the herbs Epilobium angustifolium, Maianthemum bifolium, Solidago virgaurea, Geranium sylvaticum and Trientalis europaea were sampled in early summer and late autumn from plots fertilised with a complete mixture of nutrients and from non-fertilised control plots. Concentrations of total nitrogen, insoluble and soluble proteins, free amino acids and nitrate were measured, and changes in absolute and relative concentrations of these N fractions between early summer and late autumn were used to identify the forms in which the plants store N. In all species studied, the concentration of free amino acids increased both between summer and autumn and in response to fertilisation, while the concentration of protein N increased only in response to fertilisation. Thus, free amino acids appear to have a central role in N storage. In all of the species except G. dryopteris, D. flexuosa and S. virgaurea, arginine dominated the pool of free amino acids and thus arginine was the major form of stored N in most species. In D. flexuosa and S. virgurea, however, asparagine and arginine together were the major forms of stored N, while glutamine was the major free amino acid, and N storage form, in G. dryopteris. Received: 10 March 1996 / Accepted: 22 December 1996     

影响天然林下层植物物种多样性的林分因子的研究

以我国东北过伐林区3种典型森林类型为对象,用多元逐步回归方法来研究影响天然林下层植物多样性的林分因子,尤其是与森林经营有关的林分因子,从而通过经营来维持和增加物种多样性。研究表明,与经营有关的因子中对下层植物多样性有显著影响的有林分郁闭度、公顷株数和树种多样性。因此,在林分发展的一定阶段,应采取合理经营措施如采伐来控制林分密度,保持多树种的混交,来维持和增加树种多样性。     

Effects of fire intensity on plant species composition of sandstone communities in the Sydney region

Abstract Fire intensity measures the heat output of a fire, and variation in fire intensity has been shown to have many effects on the demography of plant species, although the consequent effects on the floristic composition of communities have rarely been quantified. The effects of variation in fire intensity on the floristic composition of dry sclerophyll vegetation with different fire histories near Sydney was estimated. In particular, differences in species abundance of woodland and shrubland communities subjected to four fire‐intensity classes: unburnt, low intensity (<500 kW m^?1), medium intensity (500–2500 kW m^?1) and high intensity (>2500 kW m^?1) were examined. The samples had a standardized previous fire frequency and season, thus minimizing the effects of other aspects of the fire regime. There was a clear effect of fire intensity on the relative abundances of the vascular plant species, with increasing intensity of the fire producing vegetation that was increasingly different from the unburnt vegetation. This pattern was repeated in both the woodland and shrubland vegetation types, suggesting that it was not an artefact of the experimental conditions. However, the effects of fire intensity on floristic composition were no greater than were the differences between these two similar vegetation types, with variation in fire intensity accounting for only approximately 10% of the floristic variation. Nevertheless, the effects of fire intensity on the abundance of individual species were consistent across taxonomic groups, with the monocotyledon and Fabaceae species being more abundant at higher than lower intensities, the Proteaceae and Rutaceae more abundant at intermediate intensities, and the Epacridaceae more abundant at lower rather than higher intensities. The number of fire‐tolerant species increased with increasing fire intensity, and those fire‐tolerant species present were most abundant in the areas burnt with medium intensity. The number of fire‐sensitive species did not respond to fire intensity, and those species present were most abundant in the areas burnt with low intensity. This suggests that either fire‐sensitive species respond poorly to higher fire intensities or fire‐tolerant species respond poorly to lower fire intensities, perhaps because of differences in seed germination, seedling survival or competition among adults.     

Changes in understorey plant species richness following logging and prescribed burning in shrubby dry sclerophyll forests of south‐eastern Australia

Abstract The impacts of prescribed burning and timber harvesting on species diversity have been the subject of considerable debate. The temporal and spatial scale of these disturbances often presents major limitations to many studies. Here we present the medium‐term results of a planned long‐term study examining the impacts of logging and prescribed burning on the understorey floristic richness in shrubby dry sclerophyll forest in the south‐east of New South Wales, Australia. Generalized estimating equations were used to model the effect of environmental factors and disturbance variables on species richness at the coupe (～30 ha) and plot (～0.01 ha) scale. At the plot scale, fire effects on separate components of the vegetation were broadly consistent with other studies, with frequent fire resulting in a relative increase of species richness for species less than 1 m in height and a decline of larger species taller than this height. At the coupe scale, there was no decline in richness of larger shrub species, possibly owing to the spatial heterogeneity of fire frequency at this scale. Logging resulted in significantly greater species richness in the shrub layer, but had no significant effect on species richness in the ground layer. During the study period, there was a general decline in plant species richness at both coupe and plot scales which occurred independently of imposed management regimes. This is thought to be related to a natural succession following wildfire, and may be due to the absence of high‐intensity fire in the study area since 1973, or to an effect related to season of burning.     

Effects of tree species composition on within‐forest distribution of understorey species

Question: Do tree species, with different litter qualities, affect the within‐forest distribution of forest understorey species on intermediate to base‐rich soils? Since habitat loss and fragmentation have caused ancient forest species to decline, those species are the main focus of this study. Location: Three ancient forests, along a soil gradient from acidification‐sensitive to base‐rich, were studied: Limbrichterbosch and Savelsbos in The Netherlands and Holtkrat in Denmark. Methods: Canopy and soil surveys along transects generated data for Redundancy Analysis on tree – humus relationships. We analysed the distribution of forest plant species with Canonical Correspondence Analysis. The explanatory factors were soil characteristics (pH, organic matter, loam content and thickness of the humus layers), external crown projection, ground water and canopy data. We further analysed the relationship between forest species and humus characteristics with Spearman correlations. Results: Tree species have a significant impact on humus characteristics through the nature of their litter. Humus characteristics significantly explain the distribution of forest understorey species. The pH of the first 25 cm mineral soil and the thickness of the F‐ (fermentation) layer are the primary factors affecting the distribution of ancient forest species. Conclusion: This study indicates that the species composition of the forest canopy affects the distribution of forest understorey species. Ancient forest species are more abundant and frequent underneath trees with base‐rich litter. On acidification‐sensitive soils these relationships were stronger than on more base‐rich, loamy soils.     

连栽对桉树人工林下物种多样性的影响

采用时空序列和定位监测相结合的方法,对广西国有东门林场桉树人工林的物种多样性及其维持机制进行了研究.结果表明,桉树连栽导致人工林植物多样性减少,在667 m²样方内,第2代林的植物种类比第1代林减少了54.43%;对18块4 m²样方监测(1998～2003年)结果,第2代林植物多样性比第1代林减少了50%,物种丰富度和Shannon-Wiener指数分别比第1代林减少39.39%和17.76%.桉树人工林连栽导致群落物种多样性降低,改变了群落的物种组成及特征.不同连栽代数群落的生活型谱存在明显差异,第1代林以小高位芽、藤本高位芽和矮高位芽植物为主,而第2代林则以草本高位芽植物、地上芽植物为多;第1代林的k-对策种和r-对策种分别为69.32%和30.68%,第2代林则分别是47.83%和52.17%,可见连栽使草本植物、地上芽植物和r-对策种的数量增加,木本植物和k-对策种的数量减少.可以认为,土壤中种子库和营养体对物种多样性维持有重要作用,提出了桉树人工林群落物种多样性维持机制的初始植物繁殖体组成假说.在一个经营周期内(6～7年),桉树人工林维持的植物多样性的高低取决于初始植物繁殖体组成的丰富程度,高强度干扰的连栽方式使某些高竞争力的物种或仅以种子(果实)传播的物种丧失,造成下一代林地初始植物繁殖体组成多样性的降低,从而使连栽林地的物种多样性逐代减少.     

Effects of flooding and artificial burning disturbances on plant species composition in a downstream riverside floodplain

Our objective was to detect environmental and disturbance factors that determine plant species composition in a downstream riverside floodplain of the Shinano River, where both natural flooding and artificial fire disturbances predominate. We classified the natural vegetation into five types by physiognomy, i.e., burnt field, intact (unburnt) field, early-successional forest, later-successional forest, and margin of channel. We deployed 14 transect belts containing 713 plots of 1 m × 1 m at these sites. In the plots, we determined plant occurrences and values of five environmental (soil moisture, soil texture, relative elevation above the river, horizontal distance from the river, and light conditions) and two disturbance factors (scouring of plants by flooding and fire). Plant species compositions and environmental and disturbance factors were different among the sites. Logistic regression analysis showed that burning stimulated the occurrences of herbaceous annuals and conversely constrained those of woody and fern species, suggesting that fire disturbance resets the succession. Scouring of plants by flooding also affected the occurrences of many species, and had a positive influence on that of herbaceous winter annuals in particular. Canonical correspondence analysis indicated that the distributional characteristics of the plant species in the riverside floodplain were influenced primarily by distance from the river, reflecting the magnitude of flood disturbance. We conclude that (1) magnitude of flood disturbance is the primary factor determining plant species composition, and (2) burning maintains early successional vegetation and simultaneously creates a unique plant species composition by stimulating the germination of buried viable seeds transported by flood.     

Patterns of plant species composition on Amazonian sandstone outcrops in Colombia

Vascular plant species compositional patterns of the low forest, scrub, and herbaceous vegetation on white sand soils and sandstone substrates were studied at six sandstone plateaus in Colombian Amazonia, by means of a field survey according to the Braun‐Blanquet relevé method. Canonical Correspondence Analysis (CCA) was applied to separate effects of habitat and spatial configurations of the plateaus on species patterns. Also, information on dispersal ability and phytogeographic affinity of species was used to test explanations for between plateau differences. Low trees, shrubs and herbs were the main species recorded in 212 relevés. The main gradients in the species patterns were linked to the spatial configuration of the plateaus. Spatially controlled species patterns were mainly related to soil depth and soil organic matter. The association between phytogeographic affinity and the habitat controlled spatial link of species pointed at insufficient sampling at one plateau. Dispersal ability did not explain the habitat controlled spatially distributed occurrences of plant species. This might indicate a low frequency of local extinctions at the sandstone plateaus, especially of the poorly dispersed species, possibly because plant populations survive fire or drought disturbances in local sheltered places. Space and habitat controlled species patterns at one plateau were quite distinct from patterns at the other plateaus. This might be due to unmeasured habitat factors (e.g. unrecorded soil variation or human disturbance history) or the preferential, surveyor biased sampling procedure.     

Effects of mycorrhizal fungi on plant populations

We discuss four potentially important interactions between mycorrhizal fungi and populations of plants. First, vesicular-arbuscular mycorrhizal colonization has been shown to increase reproduction (via both male and female functions) and offspring survival, and thus it can increase population size, at least in the short term. This is undoubtedly important to wild plant species and especially to those whose success depends on high rates of reproduction such as early successional annuals. Second, the positive response in growth and reproduction to vesicular-arbuscular mycorrhizal colonization may be inversely related to plant population density. All else being equal, this would tend to stabilize the density of natural plant populations over time. It may also explain why positive responses to mycorrhizal inoculation of dense crops are rare. Third, vesicular-arbuscular mycorrhizal fungi can increase inequality in size and reproduction among plants within a population. Mycorrhizal fungi may thus exaggerate the genetic overrepresentation in the next generation of the most robust individuals in the current generation. Fourth, established mycorrhizal plants may serve as important sources of inoculum for initially nonmycorrhizal, conspecific seedlings. This may affect regeneration, and could contribute to patchy distributions of species within the community.     

Potential impacts of climate change on patterns of insect herbivory on understorey plant species: A transplant experiment

Species‐specific responses to climate change will lead to changes in species interactions across multiple trophic levels. Interactions between plants and their insect herbivores, in particular, may become increasingly disrupted if mobile herbivores respond more rapidly to climatic change than their associated host plants. We present a multispecies transplant experiment aimed at assessing potential climatic impacts on patterns of leaf herbivory. Four shrubby understorey plant species were transplanted outside their native range into a climate 2.5°C warmer in annual mean temperature. After 12 months, we assessed the types and amount of herbivore leaf damage, compared with plants transplanted to a control site within their native range. The overall amount of foliage loss to herbivores ranged from approximately 3–10% across species and sites, a range consistent with most estimates of leaf loss in other studies. The most common types of leaf damage were sucking and chewing and this pattern was consistent for all four plant species at all sites. There were no significant differences in levels and patterns of herbivory between control and warm sites for three out of four plant species. This suggests that with moderate climate warming, most herbivory will continue to be dominated by chewers and suckers, and that the overall level of foliage loss will be similar to that experienced presently.     

Land-use history affects understorey plant species distributions in a large temperate-forest complex,Denmark

In Europe, forests have been strongly influenced by human land-use for millennia. Here, we studied the importance of anthropogenic historical factors as determinants of understorey species distributions in a 967 ha Danish forest complex using 156 randomly placed 100-m² plots, 15 environmental, 9 spatial, and 5 historical variables, and principal components analysis (PCA), redundancy analysis (RDA) as well as indicator species analysis. The historical variables were status as ancient (1805 AD) high forest, reclaimed bogs, ≤100 m from Bronze Age burial mounds, or former conifer plantation, and stand age. The PCA results showed that the main gradients in species composition were strongly related to the explanatory variables. Forward variable selection and variation partitioning using RDA showed that although modern environment was the dominant driver of species composition, anthropogenic historical factors were also important. The pure historical variation fraction constituted 13% of the variation explained. The RDA results showed that ancient-forest status and, secondarily, reclaimed bog status were the only significant historical variables. Many typical forest interior species, with poor dispersal and a strong literature record as ancient-forest species, were still concentrated in areas that were high forest in 1805. Among the younger forests, there were clear floristic differences between those on reclaimed bogs and those not. Apparently remnant populations of wet-soil plants were still present in the reclaimed bog areas. Our results emphasize the importance of historical factors for understanding modern vegetation patterns in forested landscapes.     

Pollen-mediated gene dispersal within continuous and fragmented populations of a forest understorey species, Trillium cuneatum

Pollen movement plays a critical role in the distribution of genetic variation within and among plant populations. Direct measures of pollen movement in the large, continuous populations that characterize many herbaceous plant species are often technically difficult and biologically unreliable. Here, we studied contemporary pollen movement in four large populations of Trillium cuneatum. Three populations, located in the Georgia Piedmont, are exposed to strong anthropogenic disturbances, while the fourth population, located in the Southern Appalachian Mountains, is relatively undisturbed. Using the recently developed TwoGener analysis, we extracted estimates of the effective number of pollen donors (N(ep)), effective mating neighbourhood size (A(ep)) and the average distance of pollen movement (delta) for each population. We extended the TwoGener method by developing inference on the paternal gametic contribution to the embryo in situations where offspring genotypes are inferred from seeds and elaiosomes of species with bisporic megagametogenesis. Our estimates indicate that maternal plants do not sample pollen randomly from a global pool; rather, pollen movement in all four populations is highly restricted. Although the effective number of pollen donors per maternal plant is low (1.22-1.66) and pollen movement is highly localized in all populations, N(ep) in the disturbed Piedmont populations is higher and there is more pollen movement than in the mountains. The distance pollen moves is greater in disturbed sites and fragmented populations, possibly due to edge effects in Trillium habitats.     

Effects of plant species on nutrient cycling

Plant species create positive feedbacks to patterns of nutrient cycling in natural ecosystems. For example, in nutrient-poor ecosystems, plants grow slowly, use nutrients efficiently and produce poor-quality litter that decomposes slowly and deters herbivores. /n contrast, plant species from nutrient-rich ecosystems grow rapidly, produce readily degradable litter and sustain high rates of herbivory, further enhancing rates of nutrient cycling. Plants may also create positive feedbacks to nutrient cycling because of species' differences in carbon deposition and competition with microbes for nutrients in the rhizosphere. New research is showing that species' effects can be as or more important than abiotic factors, such as climate, in controlling ecosystem fertility.     

Effects of fire frequency on plant species composition of sandstone communities in the Sydney region: Combinations of inter-fire intervals

Abstract Fire frequency is the number of fires experienced by a particular community within a given time period. This variable can potentially be resolved into a number of interacting components, including time since the most recent fire, and the length of the inter-fire intervals. We estimated the effects of inter-fire intervals and sequences of inter-fire intervals, independently of time-since-fire, on the floristic composition of 26 samples from dry sclerophyll vegetation with different fire histories in Brisbane Water National Park near Sydney. In particular, we examined the effects in the recent fire history of: the length of the shortest inter-fire interval (1–26 years); repetition of very short (1–3 years) and medium (7–14 years) inter-fire intervals; and recovery from very short (1–3 years) and short (4–6 years) inter-fire intervals. Our analyses suggest that inter-fire interval may account for about 55% of the floristic variation among our samples, which were taken at a uniform 18 months after the most recent fire. There appear to be at least three general and unrelated effects of the recent history of inter-fire intervals on the floristic composition of this vegetation: (i) shorter inter-fire intervals were associated with a reduction in the number of species present in the community, along with a more unequal abundance of the remaining species, particularly of those large Proteaceae shrubs that often dominate the community biomass in dry sclerophyll shrublands of southeastern Australia; (ii) the presence of an inter-fire interval that is 1–6 years long was associated with an additional reversible reduction in the number of fire-sensitive species present in the community, particularly those common shrub species with a canopy-stored seed-bank and non-leguminous species with a soil-stored seed-bank; and (iii) repetition of inter-fire intervals that are 1–5 years long was associated with an increase in the abundance of herbaceous fire-tolerant species. This means that variation in these characteristics is associated with changes in abundance of different plant species, and that the floristic composition of any one area can be influenced in three different ways by these variables. Management of fire for plant species conservation must therefore be based on both length of time-since-fire and length of inter-fire intervals if it is to be effective.     

Effects of fire frequency on plant species composition of sandstone communities in the Sydney region: Inter-fire interval and time-since-fire

Abstract Fire frequency is the number of fires experienced by a particular community within a given time period. This concept can potentially be resolved into a number of interacting variables, including: time since the most recent fire, the length of the inter-fire intervals, and the variability of the length of the inter-fire intervals. We estimated the effects of these three variables on the floristic composition of 65 samples from dry sclerophyll vegetation with different fire histories in Brisbane Water, Ku-ring-gai Chase and Royal National Parks near Sydney. Our analyses suggest that fire frequency may account for about 60% of the floristic variation among our samples. They confirm the hypothesis that the recent (<30 years) fire frequency produces effects on floristic composition of fire-prone communities that can recognizably be attributed both to the time since the most recent fire and to the length of the intervals between fires. These effects are equal in magnitude but are different in the nature of the floristic variation they are associated with. Increasing time-since-fire is associated with a decline in the evenness of fire-tolerant species, indicating that fewer of these species come to dominate the community in the prolonged absence of fire. Herbs and small shrubs decrease in abundance, while larger shrubs increase in abundance. Inter-fire intervals of decreasing length are associated with a decrease in the evenness of the fire-sensitive species, particularly those large Proteaceae shrubs that often dominate the community biomass in dry sclerophyll shrublands of southeastern Australia. Furthermore, the variation associated with inter-fire intervals is not necessarily solely related to the shortest inter-fire interval, but is related to combinations of inter-fire intervals through time. Thus, increasing variability of the length of the inter-fire intervals is associated with an increase in the species richness of both fire-sensitive and fire-tolerant species, implying that it may be variation of the inter-fire intervals through time that is primarily responsible for maintaining the presence of a wide variety of plant species in a particular community. Our results also suggest that the floristic variation associated with different inter-fire intervals decreases with increasing time-since-fire.     

Influence of canopy tree phenology on understorey populations of Fagus crenata

Question: Is light available for subcanopy individuals of Fagus crenata spatiotemporally heterogeneous across patches with closed canopies of different foliage phenologies and gaps? Is local abundance of Sasa influenced by the composition of the canopy layer? If so, does the Sasa layer also affect the amount of light available to small F. crenata saplings? Is variation in F. crenata population structure consistent with the hypothesis that light is important? Location: Mt. Kurikoma, Japan 780 m a.s.l. Methods: Population structure of subcanopy individuals of Fagus crenata and importance of Sasa were examined in five patch types. The patch‐types were Fc (F. crenata only in the crown), Qm (Quercus mongolica var. grosseserrata only in the crown), Mo (Magnolia obovata only in the crown), Fc’ (periphery of F. crenata) and Gap. Seasonal changes in light availability above and below the Sasa layer was examined by using hemispherical photographs and quantum sensors. Results: Subcanopy individuals of F. crenata began unfolding their leaves approximately one month earlier than canopy trees of Q. mongolica var. grosseserrata and M. obovata, but a few days later than those of adult F. crenata. Accumulated photosynthetic photon flux density above the Sasa layer was greatest in Qm and Mo, and lowest in Fc. Importance of Sasa was highest in Gap. Maximum height and the number of subcanopy individuals of F. crenata were greatest in Qm, followed by Mo, and lowest in Fc. Conclusions: Differences in canopy layer composition probably influence the regeneration of F. crenata both directly through their foliage phenologies, and also indirectly by determining the importance of Sasa.     

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH₄⁺ availability in the O-horizon. We observed only short-term effects (for NO₃⁻, PO₄³⁻, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.