Banksia ericifolia transgresses the self-thinning boundary

Self-thinning B. ericifolia stands were observed to have high mean plant weights for their density, exceeding the upper boundary of the (log mean plant weight = 4.4–1.5 log mean plant density) line of White (1980) by approximately 0.5 units on the y scale (log mean weight per plant (g)). While proportion of shoot as stem and leaf area indices were similar in thinning stands of B. ericifolia and other species, specific leaf area was six to eight times less in the Banksia populations.     

Computing competition for light in the GREENLAB model of plant growth: a contribution to the study of the effects of density on resource acquisition and architectural development

BACKGROUND AND AIMS: The dynamical system of plant growth GREENLAB was originally developed for individual plants, without explicitly taking into account interplant competition for light. Inspired by the competition models developed in the context of forest science for mono-specific stands, we propose to adapt the method of crown projection onto the x-y plane to GREENLAB, in order to study the effects of density on resource acquisition and on architectural development. METHODS: The empirical production equation of GREENLAB is extrapolated to stands by computing the exposed photosynthetic foliage area of each plant. The computation is based on the combination of Poisson models of leaf distribution for all the neighbouring plants whose crown projection surfaces overlap. To study the effects of density on architectural development, we link the proposed competition model to the model of interaction between functional growth and structural development introduced by Mathieu (2006, PhD Thesis, Ecole Centrale de Paris, France). KEY RESULTS AND CONCLUSIONS: The model is applied to mono-specific field crops and forest stands. For high-density crops at full cover, the model is shown to be equivalent to the classical equation of field crop production (Howell and Musick, 1985, in Les besoins en eau des cultures; Paris: INRA Editions). However, our method is more accurate at the early stages of growth (before cover) or in the case of intermediate densities. It may potentially account for local effects, such as uneven spacing, variation in the time of plant emergence or variation in seed biomass. The application of the model to trees illustrates the expression of plant plasticity in response to competition for light. Density strongly impacts on tree architectural development through interactions with the source-sink balances during growth. The effects of density on tree height and radial growth that are commonly observed in real stands appear as emerging properties of the model.     

Native tree regeneration in native tree plantations: understanding the contribution of Araucaria angustifolia to biodiversity conservation in the threatened Atlantic Forest in Argentina

Deforestation is a global process that has strongly affected the Atlantic Forest in South America, which has been recognised as a threatened biodiversity hotspot. An important proportion of deforested areas were converted to forest plantations. Araucaria angustifolia is a native tree to the Atlantic Forest, which has been largely exploited for wood production and is currently cultivated in commercial plantations. An important question is to what extent such native tree plantations can be managed to reduce biodiversity loss in a highly diverse and vulnerable forest region . We evaluated the effect of stand age, stand basal area, as a measure of stand density, and time since last logging on the density and richness of native tree regeneration in planted araucaria stands that were successively logged over 60 years, as well as the differences between successional groups in the response of plant density to stand variables. We also compared native tree species richness in planted araucaria stands to neighbouring native forest. Species richness was 71 in the planted stands (27 ha sampled) and 82 in native forest (18 ha sampled) which approximate the range of variation in species richness found in the native forests of the study area. The total abundance and species richness of native trees increased with stand age and time since last logging, but ecological groups differed in their response to such variables. Early secondary trees increased in abundance with stand age 3–8 times faster than climax or late secondary trees. Thus, the change in species composition is expected to continue for a long term. The difference in species richness between native forest and planted stands might be mainly explained by the difference in plant density. Therefore, species richness in plantations can contribute to local native tree diversity if practices that increase native tree density are implemented.     

Competition between two naturalized dandelions,Taraxacum officinale weber andTaraxacum laevigatum DC., in mixed cultures with different levels of soil moisture

Taraxacum officinala andTaraxacum laevigatum were grown in mixed stands at various plant densities and mixing ratios with various levels of soil moisture to formulate the effect of soil moisture on the competitive relationship between the species. In pure stands, the mean plant weight—plant density relation for each level of soil moisture could be described by the reciprocal equation of the crowding effect. On the other hand, the response of mean plant weight to soil moisture content followed the reciprocal equation for a repulsive growth factor at the respective levels of plant density. By introducing the density conversion factor, the results of mixed stands could be successfully formulated from similar reciprocal equations. The dependence of density conversion factor on soil moisture content was also formulated. From these relations, a comprehensive formula was developed to describe the effects of plant density and soil moisture content on the growth of two species in mixed stands. Changes in the biomass in mixed stands were, examined by means of calculations based on the experimental results.     

Modelling the effect of autotoxicity on density-dependent phytotoxicity

An established method to separate resource competition from chemical interference is cultivation of monospecific, even-aged stands. The stands grow at several densities and they are exposed to homogenously spread toxins. Hence, the dose received by individual plants is inversely related to stand density. This results in distinguishable alterations in dose-response slopes. The method is often recommended in ecological studies of allelopathy. However, many plant species are known to release autotoxic compounds. Often, the probability of autotoxicity increases as sowing density increases. Despite this, the possibility of autotoxicity is ignored when experiments including monospecific stands are designed and when their results are evaluated. In this paper, I model mathematically how autotoxicity changes the outcome of dose-response slopes as different densities of monospecific stands are grown on homogenously phytotoxic substrata. Several ecologically reasonable relations between plant density and autotoxin exposure are considered over a range of parameter values, and similarities between different relations are searched for. The models indicate that autotoxicity affects the outcome of density-dependent dose-response experiments. Autotoxicity seems to abolish the effects of other phytochemicals in certain cases, while it may augment them in other cases. Autotoxicity may alter the outcome of tests using the method of monospecific stands even if the dose of autotoxic compounds per plant is a fraction of the dose of non-autotoxic phytochemicals with similar allelopathic potential. Data from the literature support these conclusions. A faulty null hypothesis may be accepted if the autotoxic potential of a test species is overlooked in density-response experiments. On the contrary, if test species are known to be non-autotoxic, the method of monospecific stands does not need fine-tuning. The results also suggest that the possibility of autotoxicity should be investigated in many density-response bioassays that are made with even-aged plants, and that measure plant growth or germination.     

An Individual Stand Growth Model for Mean Plant Size Based on the Rule of Self-thinning

A growth model for pure, even-aged stands of plants is asymptoticallybounded above by the self-thinning rule that relates maximumplant size to stand density. The model characterizes accretionin mean size as a deviation from the limiting size. It consistsof a function relating mean size to time and density and a companionsurvival model. The growth model is obtained by substitutingthe survival model for density in the mean size relationship.Model flexibility is demonstrated by fitting it to annual remeasurementsof mean size and number of plants per unit area in a stand ofPinus taeda L. 3/2-power rule, mortality, survival, stand dynamics, plant growth model, loblolly pine     

Mapping and characterization of mangrove plant communities in Hong Kong

Ecological surveys were carried out to investigate the distribution and characterization of remaining mangrove stands in Hong Kong. The field studies indicate that 43 mangrove stands, excluding Mai Po Nature Reserve, still remained along the coastline of Hong Kong despite tremendous reclamation and development which occurred in the past 40 years. Most mangrove stands were found in Deep Bay (western part)and Sai Kung District (eastern coasts). The total areas occupied by these mangrove stands were 178 ha,varying from a very small stand (with 1–2 mangrove shrubs) to fairly extensive mangroves in Deep Bay (> 10 ha). It appeared that mangrove stands located in Deep Bay area were larger than those in the eastern coasts. Twenty plant species were identified from these stands, with 13 being exclusive or associate mangrove species. The major constituent species were Kandelia candel, Aegiceras corniculatum, Excoecaria agallocha and Avicennia marina. Rare species such as Heritiera littoralis were only found in a few mangrove stands. Out of the 43remaining mangrove stands, 23 were more worthwhile for conservation and their plant community structures were further investigated by transect and quadrat analyses. The importance values (sum of relative abundance,frequency and dominance) show that K. candel was the most dominant species. Species richness and Simpson's indices together with tree height, tree density and canopy area fluctuated significantly between mangrove stands. These values were used to prioritize the conservation potential of the remaining mangrove stands in Hong Kong. This revised version was published online in August 2006 with corrections to the Cover Date.     

<Emphasis Type="Italic">Melaleuca quinquenervia</Emphasis> dominated forests in Florida: analyses of natural-enemy impacts on stand dynamics

Melaleuca quinquenervia (melaleuca) is a native of Australia but has become an invasive plant in Florida, USA. We conducted a long-term demographic study of melaleuca in three sections (central, transitional, and peripheral) of monoculture stands located in Florida, and quantified absolute density, diameter at breast height and basal area of trees by section at three sites. Additionally, we monitored the impacts of natural enemy (insects and fungi) on melaleuca populations which became apparent after 2001. Both absolute density and basal area, from before (1997–2001) and after noticeable natural-enemy impact (2002–2005), were compared. Prior to the natural-enemy impact, absolute density of melaleuca trees declined primarily due to self-thinning and associated losses of small trees, but diameter at breast height increased, as did the basal area. Later during the period when natural enemies prevailed, absolute density declined at a significantly greater rate across all sections but was highest at the periphery. The decrease in mean absolute density and basal area/ha of melaleuca during the natural-enemy impacted period coincided with the increased incidence of the populations of plant-feeding insects and fungi. The mean diameter at breast height continued to increase in all sections of the stands throughout the study period. An increasing trend in basal area prior to natural-enemy impact was reversed after increase in natural-enemy abundance and noticeable impact in all three sections of the stands. These findings lend support to a growing body of literature that implicates natural enemies as increasingly important density-independent regulators of M. quinquenervia populations. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged     

Effects of longleaf pine planting density and other factors on stand structure and associated wildlife habitat

The primary objective of many longleaf pine (Pinus palustris) restoration programs is to enhance or restore habitat for wildlife dependent on herbaceous plant communities. Because herbaceous cover is inversely related to canopy cover, restoration programs often place restrictions on longleaf pine planting density. However, the influence of planting density on understory plant communities has been inadequately evaluated. Therefore, we initiated a study to examine the relative influences of planting density and other factors on overall understory composition and forage availability for white‐tailed deer (Odocoileus virginianus) and northern bobwhite (Colinus virginianus) in nine longleaf pine stands throughout the Coastal Plain of Alabama during 2017–2018. We found that coverage of herbaceous plants decreased 3.5%, coverage of woody plants decreased 2.4%, and coverage of northern bobwhite forage plants decreased 1.9% for each 1 m²/ha increase in longleaf pine basal area. However, planting density was not a significant predictor of current basal area, nor coverage of any functional group of plants we examined, likely because current longleaf pine density averaged only 46% (range = 30–64%) of seedling planting density. We did not detect an effect of prescribed fire on stand condition or understory plant communities, likely due to variability in fire timing and frequency. Our findings related to planting density were likely a function of low longleaf pine survival, which is not uncommon. Because of this and the inherent variability in growth rates for young longleaf pine stands, restoration programs should consider placing greater emphasis on post‐planting monitoring and management than planting density.     

Uncommon vascular plant species in an East-Central Swedish forest area-a comparison between young and old stands

Abundance of uncommon vascular plant species (indicator species and red-listed species) was recorded in 120 circular (12.6 m²) random plots in young (10–24 years after deal-cutting) and old (>80 years) forest stands, respectively, in the forest area of Vällen (2 600 ha), ca. 80 km north of Stockholm, Sweden. Factors describing stand density, field-layer and soil moisture were recorded. A total of 16 species were observed and of these 14 were found in old and 9 in young stands. Seven species occurred exclusively in the old stands and two in the young stands. The total number of species records was significantly higher in young than in old stands, 80 and 71, respectively. Mean number of records per ha was ca. 530 for the young stands and ca. 470 for the old. Of the five species common enough to be tested, a difference was only found for Paris quadrifolia , which was significantly more frequent in young than old stands. Galium odoratum, Hepatica nobilis, Lathyrus vernus and Paris quadrifolia grew in significantly more sheltered sites, e.g. beneath other herbs and grasses or close to shrubs and trees in young compared to old stands. Nested subset analysis showed that the whole data-set had a nested structure. Galium odoratum was significantly nested and thus indicated rich plant assemblages. In this calcareous area with nutrient-rich forest soils, there evidently is a potential for at least some uncommon vascular plant species to survive the phases following clearcutting. The study indicates that green-tree retention at final fellings and avoidance of heavy shrub removal during cleaning are beneficial to the flora.     

Influence of Post‐Clearing Treatment on the Recovery of Herbaceous Plant Communities in Amazonian Secondary Forests

Secondary forests are an increasingly common feature in tropical landscapes worldwide and understanding their regeneration is necessary to design effective restoration strategies. It has previously been shown that the woody species community in secondary forests can follow different successional pathways according to the nature of past human activities in the area, yet little is known about patterns of herbaceous species diversity in secondary forests with different histories of land use. We compared the diversity and abundance of herbaceous plant communities in two types of Central Amazonian secondary forests—those regenerating on pastures created by felling and burning trees and those where trees were felled only. We also tested if plant density and species richness in secondary forests are related to proximity to primary forest. In comparison with primary forest sites, forests regenerating on non‐burned habitats had lower herbaceous plant density and species richness than those on burned ones. However, species composition and abundance in non‐burned stands were more similar to those of primary forest, whereas several secondary forest specialist species were found in burned stands. In both non‐burned and burned forests, distance from the forest edge was not related to herbaceous density and species richness. Overall, our results suggest that the natural regeneration of herbaceous species in secondary tropical forests is dependent on a site's post‐clearing treatment. We recommend evaluating the land history of a site prior to developing and implementing a restoration strategy, as this will influence the biological template on which restoration efforts are overlaid.     

不同密度巨桉人工林林下植物多样性及根际土壤化感物质

为探究巨桉人工林林下植物多样性的影响因素,选择四川省丹棱县4和8年生不同密度(1200、1600和2000株·hm^-2)巨桉人工林为对象,研究林下植物多样性和根际土壤酚类化感物质。结果表明: 共发现植物45种,隶属于33科44属;随林龄增加和林分密度降低,巨桉人工林林下植物种类增加且重要值分布更为均匀;各林分植物生活型以高位芽植物为主。4年生林地灌木Shannon指数和Margalef指数在1600株·hm^-2下显著升高,4年生林地除Margalef指数外其余草本多样性指数及8年生林地草本Pielou指数随林分密度降低均显著升高;灌木Shannon指数及Margalef指数在1200株·hm^-2下8年生林地显著高于4年生林地。根际土壤中鉴定出5种酚类化感物质,4年生林地根际土壤中水杨酸浓度在1600株·hm^-2下显著降低,5,7-二羟基黄酮浓度随林分密度降低而显著降低;水杨酸浓度在8年生林地根际土壤中随林分密度降低显著升高;水杨酸浓度在2000株·hm^-2下4年生林地显著高于8年生林地,在1600株·hm^-2下相反;5,7-二羟基黄酮浓度在1200株·hm^-2下8年生林地显著高于4年生林地。冗余分析(RDA)结果表明,土壤pH、容重、有机质、全磷、全氮及化感物质是林下植物多样性的主要环境影响因子。适度延长轮伐期、调节林分密度以改善林内微环境、缓冲巨桉的化感作用,可促进林下植被发育。     

Long-term growth dynamics of natural forests in Hokkaido,northern Japan

Abstract. The long-term growth dynamics of natural forest stands on the island of Hokkaido were described on the basis of an analysis of data from 38 permanent plots spanning 15–22 yr. Stand structure was characterized by basal area, stem density and tree size variability. To detect trends in stand structure, regression models for recruitment rate (per ha per yr), mortality rate and the rate of change in stem density and tree size variability were developed by a stepwise method using initial basal area, stem density, tree size variability, species composition summarized by LNMDS ordination, altitude, annual mean temperature, annual precipitation, type of understorey vegetation, topography and slope aspect as candidates for predictor variables. The same analyses were conducted for basal area increment (net growth) and its components: survivor growth = basal area gain by growth of surviving individuals and mortality = basal area loss by death of individuals. Stem density remained generally unchanged; recruitment was relatively low even in very sparse stands. Stand basal area generally increased as survivor growth was approximately double the mortality. Recruitment rate was strongly affected by the presence of dwarf bamboo (Sasa spp.) vegetation on the forest floor which inhibited tree regeneration. Mortality rate was density-dependent; dense stands had higher mortality than sparse stands. Density change rate (recruitment rate - mortality rate) was, therefore, determined by both the type of understorey vegetation and stem density. Survivor growth was high in stands with high stem density and basal area. Mortality was dependent on basal area and altitude. Net basal area increment (net growth) was dependent only on stem density with other factors that influenced survivor growth and mortality omitted. Tree size variability decreased in stands with high tree size variability whereas it increased in stands with low size variability. Based on the obtained models for density change rate and net basal area increment, trajectories of stands were illustrated on a log-log diagram of stem density and basal area. The predicted differences in trajectories as affected by the understorey vegetation type indicated the importance of dwarf bamboo vegetation for forest dynamics on Hokkaido.     

Vegetation analysis and tree population structure of tropical wet evergreen forests in and around Namdapha National Park,northeast India

Species composition, diversity and tree population structure were studied in three stands of the tropical wet evergreen forest in and around Namdapha National Park, Arunachal Pradesh, India. Three study stands exposed to different intensities of disturbances were identified, viz., undisturbed (2.4 ha) in the core zone of the park, moderately disturbed (2.1 ha) in the periphery of the park and highly disturbed (2.7 ha) outside the park area. In total 200 plant species belonging to 73 families were recorded in three stands. Tree density and basal area showed a declining trend with the increase in disturbance intensity. The densities of tree saplings and seedlings were lower in the disturbed stands than in the undisturbed stand. Species like Altingia excelsa, Olea dioica, Terminalia chebula, Mesua ferrea and Shorea assamica in the undisturbed stand and Albizia procera alone in the moderately disturbed stand contributed more than 50% of the total tree density in respective stands. The undisturbed stand contained young tree population. In the highly disturbed stand, the tree density was scarce, but had uncut trees of higher girth class (>210 cm GBH). Low shrub density was recorded in both disturbed stands due to frequent human disturbances; the broken canopy and direct sunlight enhanced the abundance of herbs in these stands. With a species rarity (species having <2 individuals) of ca. 50%, the tropical wet evergreenforests of the Namdapha National Park and its adjacent areas warrant more protection from human intervention and also eco-development to meet the livelihood requirements of the local inhabitants in the peripheral areas of the Namdapha National Park in order to reduce the anthropogenic pressure on the natural resources of the park.     

The influence of plant density on the responses of Sinapis alba to CO2 and windspeed

Plants in nature live in populations of variable density, a characteristic which may influence individual plant responses to the environment. We investigated how the responses of Sinapis alba plants to different wind speeds and CO₂ concentrations could be modified by plant density. In our wind-density experiment the expectation that mechanical and physiological effects of wind will be ameliorated by growing in high density, as a result of positive plant interactions, was realised. Although individual plants were smaller at higher densities, the effect of increasing windspeed was much less than at lower plant densities. A similar reduced sensitivity of individual plant growth under high densities was also observed under CO₂ enrichment. When measured as a population or stand response, there was no effect of density on the CO₂ responses, with all stands showing very similar increases in total biomass with CO₂ enrichment. In the wind speed experiment, total biomass per stand increased significantly with density, although there was no effect of density on the wind speed response. Specific leaf area decreased with increasing wind speed and this response was significantly affected by the density at which the plants grew.     

Factors of pine-stand transformation in the city of Yekaterinburg

The relationships between the health of Pinus sylvestris trees and stands and the levels of anthropogenic stress and fragmentation of habitats have been studied based on the example of pine stands of natural origin in Yekaterinburg and its vicinity (48 sampling plots located at different distances from the center of the city). The distances from the city center and from the boundary of a built-up area to a sampling plot and the population density around this plot are taken to characterize the level of anthropogenic stress. The area and perimeter of a forest site; the distance from the boundary of a sampling plot to the forest edges; and the shape of forest edges are used to characterize the level of fragmentation of habitats. We have found that the average morphological metrics of trees have slight changes, while the traits of tree health are significantly altered under urban environment. The overall performance of stands decreases by 22–35% in the urbanization gradient. The health of stands is most substantially influenced by the forest-site area. The population density around sampling plots and the distance of these plots from the megalopolis center have a lower effect on the health of trees and stands. Fragmentation of forests in the city leads to a decrease in the size of forest sites, which causes a decrease in wood stocks and the diameter and height of trees and the deterioration of their health. The lowest area of the forest to make it sustain an urban environment is about 30 ha. The untransformed core of forest is located no less than 70 m from the forest edge.     

The effect of clumped planting patterns on epidemics of damping-off disease in cress seedlings

Summary Environmentally controlled experiments on damping-off disease of garden cress (Lepidium sativum) caused by Pythium irregulare were used to study the effect of clumped planting patterns upon epidemic rates. In clumped stands of seedlings the rate of multiplication of disease was, in general, slower than in unclumped, evenly spaced stands of the same overall density. When the number of clumps per unit area was varied, but the proportion of the total area occupied by clumps kept constant, there was no significant change in the rate of linear advance of the disease between different treatments, but the multiplication rate in randomly inoculated plots was higher at 100 clumps/m² than 200 clumps/m² due to an unexpected interaction between the multiplication rate and the distribution of primary disease foci. When the area occupied by clumps was varied, but the number of clumps per unit area and the overall plant density were kept constant, both the multiplication rate and the rate of linear advance of the disease were significantly reduced in the stands with the most condensed clumps.     

Shoot dynamics of the giant grass Gynerium sagittatum in Peruvian Amazon floodplains,a clonal plant that does show self-thinning

The giant rhizomatous grass Gynerium sagittatum is an early successional species that forms dense monocultures in Peruvian Amazon floodplains. We studied the shoot population structures by recording shoot densities and shoot heights. Leaf areas and stem volumes were allometrically estimated. Stands of two varieties of G. sagittatum were examined that differ in height and in the degree of shoot branching. In stands of increasing age, marked decreases in shoot densities were accompanied with an increase in mean shoot size. Self-thinning was indicated by the negative correlation between log stem volume per unit ground area and log shoot density, significant at least for one of the two varieties. The difference in thinning slope between the varieties could be largely accounted for by their different shoot geometry, as was revealed by calculations based on the allometric model of Weller (1987b). The relationship between log leaf area per shoot and log shoot density was significantly negative with slopes close to –1. Shoot size inequalities decreased with increasing mean stem volume per shoot, probably as a result of density-dependent mortality of the smaller shoots. All of these results accord with expectations for shoot self-thining. Gynerium sagittatum is the first clear example of a clonal plant species that exhibits self-thining in natural monospecific stands. It is argued that self-thinning occurs in this giant tropical grass because its shoots are perennial and do not experience seasonal die-back (periodic density-independent mortality), in contrast to many of the clonal plant species that have been studies so far.     

Structure and floristics of secondary and old-growth forest stands in lowland Costa Rica

We characterized stand structure and floristic composition of woody life forms in three, 16–18 yr old secondary stands that regenerated after pasture abandonment, and three nearby old-growth stands of tropical rain forest in lowland Costa Rica. Basal area and stem density for each of four plant size classes (seedlings, saplings, treelets, trees) were similar among stand types, but density of adult canopy palms (individuals 10 cm DBH), was lower in the secondary stands. We estimate that 15% of the basal area of stems 10 cm DBH correspond to remnant trees in our secondary stands. The observed rapid woody regrowth compared to other published studies in the lowland neotropics, can be attributed to moderate land use and possibly, to the influence of nutrient-rich volcanic soils in the study area. Overall, plant species richness was lower in the secondary stands, but this difference was less pronounced in the smallest size classes (seedlings, saplings). Median percent similarity of all pairwise stand comparisons showed that floristic composition of saplings (stems 1 m tall and 5 cm DBH) was more similar between secondary and old-growth stands than for trees (stems 10 cm DBH). Because the potential value of secondary forests in conserving woody plant diversity appears highest for the young size classes, we suggest that further studies on floristic composition, especially those addressing the dynamics of the understory component, are needed to refine our understanding of the role of this natural resource in the maintenance of plant biodiversity in disturbed landscapes.     

Interspecific competition for seed production between two naturalized dandelions under different levels of fertilizer supply

Two naturalized dandelions,Taraxacum officinale andTaraxacum laevigatum, were grown in mixed stands at various plant densities and mixing ratios with various levels of fertilizer supply until seeds were produced. The effect of fertilizer supply on the competitive relationship for seed production between the species was analyzed. In pure stands, the simultaneous effects of plant density and fertilizer supply on mean plant weight were formulated for each species. By introducing the density conversion factor, the results of plant weight growth in mixed stands were formulated successfully.T. laevigatum produced more seeds thanT. officinale at the same plant weight. The relationship between mean plant weight and mean number of mature seeds produced per plant was also formulated for each species. Ratio diagrams were predicted with these formulas. The density conversion factors showed thatT. officinale was always dominant overT. laevigatum for plant weight growth within a generation. The ratio diagrams showed thatT. officinale became dominant overT. laevigatum for seed production with repeating generations at higher levels of fertilizer supply, but that the species with a larger ratio of plant density became dominant over the other species at lower levels of fertilizer supply.