Why is Aucuba japonica Smaller in Heavy Snowfall Areas? A Growth Simulation of Evergreen Broad-leaved Shrubs based on Shoot Allometry, Critical Shoot Sizes for Flowering and Photosynthetic Production

Aucuba japonica , an evergreen broad-leaved shrub. The model is based on the allometry of plant organs and incorporates the processes of flowering, branching, and allocation of biomass among different organs. Growth of model plants was simulated under different values of annual gross production per leaf area (LGP) for 15 years. The tree form was mainly determined by the critical shoot size for initiating flowering (SF), but the average increase of new shoot mass (INM) for two successive years had marked effects on the growth pattern of A. japonica in addition to SF. Under small LGP conditions, plants with larger SF did not flower or postponed the initiation of flowering. With increasing LGP, plants with larger SF had a greater advantage because of their high efficiency for vertical growth. A smaller SF was observed in A. japonica var. borealis, which is distributed in heavy snowfall areas, and resulted in precocious flowering with a higher reproductive rate. The small plant size and shoot size of var. borealis may be the result of selection for preventing the delay of the year in which reproduction starts under smaller LGP conditions. On the other hand, the large SF of A. japonica var. japonica, which is distributed in light snowfall areas, suppresses flowering under small LGP conditions, and flowering occurs only after achieving a certain amount of vertical growth. Received 8 July 1999/ Accepted in revised form 21 December 2000     

Comparison of ecophysiological responses to heavy snow in two varieties ofAucuba japonica with different areas of distribution

Aucuba japonica varieties are common evergreen understory shrubs in Japan.Aucuba japonica var.borealis is distributed on the Sea of Japan side of Honshu and Hokkaido where heavy snow cover lasts for more than 3 months in winter.Aucuba japonica var.japonica is distributed in areas with shallow or no snow on the Pacific Ocean side of Honshu and Shikoku. The ecophysiological characteristics of var.borealis were compared with those of var.japonica to examine the effects of heavy and long-term snow cover on the life cycle of var.borealis. Shoots of both varieties were shaded in crushed ice for 110 days, but their photosynthetic activities, chlorophyll contents and the chlorophylla/b ratio was not affected. The leaves of var.borealis were no less frost tolerant than those of var.japonica. In spite of the difference in environmental factors, both varieties had similar characteristics in seasonal changes of photosynthesis, respiration and chlorophylla/b ratio. These results suggest that var.japonica could survive in areas with heavy snow where it does not normally occur. Leaf net production (LNP) was estimated based on the microclimatic data and seasonal photosynthetic and respiration rates. The difference in the annual LNP between the two varieties was equivalent to the difference in the LNP during the snow season. One of the major effects of snow cover is to interrupt and reduce the production period of var.borealis.     

Two types of matter economy for the wintering of evergreen shrubs in regions of heavy snowfall

Plant adaptation to an environment subject to heavy snowfalls was investigated in four species of evergreen shrubs growing in a Fagus crenata forest in an area of Honshu on the Sea of Japan. These shrubs stored carbohydrates in some organs before the snowy season and were covered with snow for 4–5 months. Aucuba japonica var. borealis, Camellia rusticana, and Ilex crenata var. paludosa maintained a reserve of carbohydrates during the snowy season. In Daphniphyllum macropodum var. humile, the reserve of carbohydrates decreased during winter. The respiration rates in the first three species decreased from autumn to winter, whereas the decrease in D. macropodum was slight. It was found that the first three species could use reserve carbohydrates for the growth of new shoots after the thaw, whereas in the last species the growth of new shoots depends on high photosynthetic activity in late spring. Our findings suggest some types of matter economy in evergreen shrubs for wintering in an environment of heavy snow.     

Consequences of the Elimination of Old Leaves upon Spring Phenological Events and the New Leaves Nutrient Concentration in a Wintergreen Woody Species in the Southern Hemisphere

Previous studies analyzed the importance of old leaves conservancy for wintergreen species plant growth only after early spring old leaves elimination. However, carbon and nutrient resources for growth could have already been translocated from old leaves to shoots during autumn. In this work, the effect of old leaves absence on the leaf mass per area (LMA, g m⁻²) and nutrient concentration of new spring leaves, shoot growth, and flowering was studied in Aristotelia chilensis, an Andean Patagonic woody wintergreen species of Argentina. Plants were studied after autumn defoliation (AD) or late winter defoliation (WD) and results were compared to those of undamaged control plants (CO). The new leaves LMA and mineral nutrient (N, P, K, and Mg) concentration values did not decrease in AD or WD compared to CO plants. Conversely, CO plants showed higher flowering intensity and shoot lengthening compared to AD or WD plants. There were not remarkable differences regarding the defoliation time, though non-flowering shoots grew in a lesser degree than the flowering shoots in WD plants. It was concluded that A. chilensis old leaves cohort is an important source to shoot growth and flowering but their absence does not affect the new leaves structure or nutritional status from early spring in either AD or in WD plants. New leaves formation probably is guaranteed by resources (carbon and nutrients) previously stored in stems or even in the buds containing the preformed leaves since March, by the end of summer. Provided the availability of complete resources for the new leaf flush independently of the old leaves A. chilensis would restore the carbon balance as soon as possible to resume the growth of heterotrophic tissues at normal rates. Endogenous response to counterbalance the old leaves absence on non-flowering shoots was more effective when there was greater lag time between defoliation and shoot growth resume. Flowering and non-flowering shoots compete for the available resources when A. chilensis have not yet expanded leaves and shoots supporting reproductive structures were stronger sinks compared to non-flowering shoots in WD plants.     

Carbon autonomy of reproductive shoots of Siberian alder (<Emphasis Type="Italic">Alnus hirsuta</Emphasis> var.<Emphasis Type="Italic"> sibirica</Emphasis>)

Carbon autonomy of current-year shoots in flowering, and of current-year shoots plus 1-year-old shoots (1-year-old shoot system) in fruiting of Siberian alder (Alnus hirsuta var. sibirica) was investigated using a stable isotope of carbon, ¹³C. The current-year shoot and 1-year-old shoot systems were fed ¹³CO₂ and the atom% excess of ¹³C in flowers and fruits was determined. The majority of photosynthate allocated to flower buds was originally assimilated in the leaves of the flowering current-year shoots. Of all the current-year shoots on fruiting 1-year-old shoots, only those nearest to the fruits allocated the assimilated photosynthate to fruit maturation. These results indicate that the current-year shoots and 1-year-old shoot systems are carbon-autonomous units for producing flowers and maturing fruits, respectively.     

Management of below-ground biomass of <Emphasis Type="Italic">Typha angustifolia</Emphasis> by harvesting shoots above the water surface on different summer days

A dynamic model of regrowth in Typha angustifolia after cutting shoots above the water surface was formulated by characterizing the phenology and mobilization of resources from below-ground to above-ground organs after the cutting. The model parameters were determined by two cutting experiments to investigate the different strategies with flowering and non-flowering shoots after cutting in 2001 and by four cutting experiments to elucidate the regrowth characteristics after cutting on different days from June to September in 2002. A difference was evident both for flowering and non-flowering shoots and for each cutting day. From June to August, non-flowering shoots regrew immediately after cutting, but flowering shoots did not. The shoot regrowth height, number of leaves and shoot biomass were higher with the earlier cutting. The model was validated using the below-ground biomass observed in December 2002 and below-ground dynamics observed in 2003. In the low-flowering shoot zone of the stands, in which the percentage of flowering shoots was small (around 10%), the decrease in below-ground biomass became larger from June (20%) to August (60%). Cutting the high-flowering shoot zone (flowering shoots: 78%) in July 2001, just 1 week after peduncle formation, decreased the below-ground biomass by about 50%. In the low-flowering shoot zone, cutting just before senescence is better for decreasing below-ground biomass with a smaller rate of flowering shoots. The difference of below-ground biomass reduction in non-flowering shoots is mainly due to the decrease in downward translocation (DWT) of above-ground material to below-ground organs during senescence, because of the decrease in regrowth biomass. As for flowering shoots, the decrease in the photosynthate transportation from above-ground to below-ground organs and that of DWT are closely related because they cannot grow again within the season.     

Differences in Architecture and Shoot Growth during Stagnant and Extension Growth Phases of Acanthopanax sciadophylloides(Araliaceae)

The shoot growth of a deciduous tree, Acanthopanax sciadophylloidesFranch. et Savat. shows inter-annual intermittent repetitionof two distinctive phases, a stagnant growth phase (S-phase)and vigorous extension-growth (E-phase). To help understandthe differentiation mechanism, shoot development was studiedover time in both shoot phases. S-phase and E-phase shoots weredistinguished from each other by morphological traits: S-phaseshoots are characterized by higher allocation to leaves anda shorter period of stem growth, while E-phase shoots show continuousstem extension over the growing season. Specific leaf area didnot differ between the two phases. This shoot differentiationwas similar to the morphological differentiation of shoots betweenlong vs. short shoots found in some temperate trees. Leavesof both phases were well-dispersed through adjustment of petiolelength and leaf-blade size to reduce mutual shading within ashoot. Stem-wood density of current-year shoots was lower inE-phase compared with S-phase shoots. Leaves produced earlyin the season affected the growth phase of the following year.These results suggest that annual shoot differentiation of A.sciadophylloides was determined during the previous season andreflects leaf productivity in a given habitat during that growingseason. Copyright 2001 Annals of Botany Company Acanthopanax sciadophylloides, Araliaceae, biomass allocation, intermittent shoot growth, leaf display, shoot architecture, shoot differentiation     

Population dynamics,productivity and biomass allocation ofZizania latifolia in an aquatic-terrestrial ecotone

The population and production ecology of aZizania latifolia stand at a sheltered shore of the Hitachi-Tone River were investigated. Shoot emergence was observed twice a year; the fist was a synchronized shoot emergence in April and the second was from August to October. Aboveground biomass was mostly occupied by leaves and peaked at 1500 g dry weight m⁻² in August. The belowground biomass also reached its peak, 750 g dry weight m⁻², in August. The secondary shoots were small in spite of their high density. Leaves were produced continuously throughout the season. The leaf life span was as short as 55.6 days for cohorts that emerged from May through to September. Total annual net production ofZ. latifolia could be more than 3400 g dry weight m⁻². Shoot clusters of several centimeters were observed in April. The following self-thinning caused a regular distribution of the remaining shoots in August. Most shoots produced in August to October were found near a shoot persisting since April. They showed more concentrated distribution than shoots in April. A large biomass allocation to leaves and the ability to produce many clump shoots during the late growing period may facilitate dominance ofZ. latifolia in relatively sheltered sites.     

Effects of simulated herbivory in three old field Compositae with different inflorescence architectures

The effects of simulated herbivory (early or late defoliation and cutting of the flowering shoot) on the growth and reproduction of three species of monocarpic composite forbs (Crepis pulchra, Picris hieracioides and C. foetida) with different inflorescence architectures were studied in experimental plots. For the three species studied, early defoliation had no significant effect on subsequent growth. In contrast, late defoliation, occurring at the start of the season of drought, had a negative effect on growth and reproduction in the two Crepis species, particularly C. foetida, but had less effect on P. hieracioides. Sexual biomass was more clearly affected by late defoliation than was vegetative biomass, although the effects differed markedly among species possibly as a result of differences in phenology. Clipping the flowering shoot removed about 3 times less biomass than late defoliation and had little effect on vegetative biomass. It had much greater effects on the sexual biomass in P. hieracioides and C. pulchra, and resulted in the production of many shoots sprouting from the rosette, allowing the treated plants to regain a vegetative biomass close to that of control plants. Clipping did however lead to the production of shorter shoots and a reduction in the number of capitula formed. In C. foetida, much branching occurred even when the main shoot was not cut; the architecture of individual plants was therefore only slightly changed by clipping the apical bud and the sexual biomass of this species was not affected by ablation of the flowering shoot. Overcompensation was found in only two families of C. pulchra for vegetative biomass. No over-compensation was found for sexual biomass, despite an increase in the number of flowering shoots in C. pulchra and P. hieracioides following clipping. However situations close to compensation for the vegetative biomass in the three species and in P. hieracioides for the sexual biomass were recorded. The response of the three study species to simulated herbivory were related to their architecture and to the time of defoliation.     

Shoot growth dynamics and size-dependent shoot fate of a clonal plant,Festuca rubra, in a mountain grassland

The relation of the within-season and between-season patterns of shoot growth were compared in a clonal grass with long-lived shoots,Festuca rubra, in a mown mountain grassland. The growth rate of shoot length from spring to summer in a year was almost constant for each shoot irrespective of spring shoot length each year. The annual shoot growth rate from spring to spring was negatively correlated with the shoot length in the first spring. Shoots of different length and age therefore tended to converge over time to a population of identical shoot size, suggesting an equalizing effect of growth pattern on size structure. Shoot size (shoot length and number of leaves) influenced the fates of shoots. Larger shoots showed an increased incidence of both flowering and formation of intravaginal daughter shoots and a decreased incidence of death in the subsequent time period. The fates of shoots were independent of their age. Although the negatively size-dependent springto-spring annual shoot growth rate acted to decrease shoot size variation, the remaining variation within the shoot population was still sufficient to generate different fates of shoots. These fates were not related to the previous life history of individual shoots. There was a significantly positive effect of the shoot size at initiation on its life expectancy. This was mainly attributable to the positively size-dependent survival rate of shoots in the early stage (<1 year old) of shoot life history. Later on (> 1 year old), shoot size had little effect on the survival rate of shoots. Once small young shoots have survived this early stage (< 1 year old) in life history, they can grow vigorously, little affected by competition regardless of shoot size, and converge to a stable size structure of shoots of similar size. Only shoot size in the early stage ( < 1 year old) of life history is important for the persistence of a shoot population.     

Flower bud abortion influences clonal growth and sexual dimorphism in the understorey dioecious shrub Aucuba japonica (Cornaceae)

Sexual differences were investigated to determine the significance of flower bud abortion in the dioecious shrub Aucuba japonica Thunb. The mean number of flowers per inflorescence and the mean number of flowering inflorescences (as opposed to aborted inflorescences) per individual were greater in males than in females in 1997 and 1998. Reproductive investment by males was 0.4-times (1997) and 1.4-times (1998) that by females. In addition, females aborted 30.9% (1997) and 42.7% (1998) of their total flower buds without blooming, whereas no male flower buds aborted. One of the architectural traits of this shrub is that in the year that a flower bud is produced at the shoot apex, the shoot will branch into two or more shoots. Thus, there was less sexual difference in the number of current shoots per individual than there was in the number of flowering inflorescences. The relationship between annual growth and reproduction, and the probability of reproduction in the following year, suggested that the higher investment in female reproduction was manifested as a cost for reproductive frequency rather than as a cost for annual growth. The spatial distribution of both males and females was clumped, which may be the result of clonal growth. In addition, overall sex ratios were not skewed and the number of sprouts did not differ significantly between sexes. These results suggested that flower bud abortion by females might reduce sexual dimorphism in terms of clonal growth.     

Structural cost of shoot modules and its implications on plant potential fitness in a Mediterranean perennial shrub, Retama sphaerocarpa (L.) Boiss

This paper introduces a methodology to analyse the structural costs on plant potential fitness, empirically exemplified in the hierarchical shoot system of a Mediterranean perennial plant, Retama sphaerocarpa (L.) Boiss. During growing season every year (March-August), the terminal shoot (which is the basic unit of growth) develops inflorescences, flowers and fruits, as well as new shoots (first-, second- and third-order branching shoots) which have the potential to "behave" as terminal shoots in the following year. Different morphological and demographical aspects of the modules within the terminal shoot were measured in 100 terminal shoots selected from different plants of a natural population of R. sphaerocarpa. Complementary samples of 100 shoots of different branching orders were collected to obtain biomass estimations of the terminal shoots. We propose a simple procedure to estimate structural cost (biomass investment) on plant potential fitness (flowering buds) as a methodology for interpreting and comparing the consequences on fitness of different plant growth patterns. The results of this study exemplify how differential allocation patterns among plant structural modules, depending on their position within the shoot system, can be quantified to estimate their influence upon plant potential fitness.     

The influence of below ground herbivory and plant competition on growth and biomass allocation of purple loosestrife

Experiments investigating plant-herbivore interactions have primarily focused on above-ground herbivory, with occasional studies evaluating the effect of below-ground herbivores on plant performance. This study investigated the growth of the wetland perennial Lythrum salicaria (purple loosestrife) under three levels of root herbivory by the weevil Hylobiustransversovittatus and three levels of plant competition by the grass Phleumpratense in a common garden. Plant growth, flowering phenology, and biomass allocation patterns of purple loosestrife were recorded for two growing seasons. During the first year, root herbivory reduced plant height; plant competition delayed flowering; and the interaction of root herbivory and plant competition resulted in reductions in plant height, shoot weight and total dry biomass. Plant competition or larval feeding did not affect the biomass allocation pattern in the first year. These results indicate the importance of interactions of plant competition and herbivory in reducing plant performance – at least during the establishment period of purple loosestrife. In the second growing season, root herbivory reduced plant height, biomass of all plant parts, delayed and shortened the flowering period, and changed the biomass allocation patterns. Plant competition delayed flowering and reduced the dry weight of fine roots. The interaction of root herbivory and plant competition delayed flowering. Root herbivory was more important than plant competition in reducing the performance of established purple loosestrife plants. This was due, in part, to intense intraspecific competition among the grass individuals effectively preventing shoot elongation of P. pratense and resulting in a carpet like growth. Received: 3 April 1997 / Accepted: 27 July 1997     

Endophytic and epiphytic phyllosphere fungi of red-osier dogwood (<Emphasis Type="Italic">Cornus stolonifera</Emphasis>) in British Columbia

Endophytic and epiphytic phyllosphere fungi associated with red-osier dogwood (Cornus stolonifera), a deciduous shrub, were examined in coastal British Columbia, Canada. Current-year shoots were divided into four types based on the absence or presence of inflorescence and secondary elongated shoots at the apex of primary shoots. Leaves on these shoots were then classified into six categories so as to examine the effect of flowering, secondary shoot elongation, and shoot order within current-year shoots on the occurrence of phyllosphere fungi. Species composition of fungi was markedly different between the interior and surface of leaves, whereas it was relatively similar among the six leaf categories in the interior or on the surface. Frequencies of the eight major species were not different between leaves on flowering and nonflowering shoots. The frequency of Colletotrichum gloeosporioides in the leaf interior was greater on leaves on the primary shoots that elongated the secondary shoots than on those that did not, and was greater on leaves on the primary shoots than on those on the secondary shoots. On the other hand, secondary shoot elongation and shoot order had no effect on the frequencies of C. gloeosporioides and the other seven epiphytes on leaf surfaces.     

Variation in shoot mortality within crowns of severely defoliated <Emphasis Type="Italic">Betula maximowicziana</Emphasis> trees in Hokkaido,northern Japan

To clarify mortality patterns of current-year shoots within the crown of Betula maximowicziana Regel after severe insect herbivory in central Hokkaido, northern Japan, we investigated the degree of defoliation, pattern of shoot development, shoot mortality, and leaf tissue-water relations. One hundred current-year long shoots growing in a B. maximowicziana plantation were observed for defoliation and mortality in June 2002. An outbreak of herbivorous insects (Caligula japonica and Lymantria dispar praeterea) occurred in the stand in mid-to-late June, and the monitored shoots were defoliated to various degrees. Within 1 month of defoliation, some of the severely defoliated shoots had produced new leaves on short shoots that had emerged from axillary buds. Stepwise logistic regression revealed that the probability that current-year long shoots would put out axillary short shoots with leaves is closely related to the degree of defoliation. To evaluate the water relations of the leaves, we determined pressure–volume curves for the leaves that survived the herbivorous insect outbreak and the new leaves that emerged after defoliation. The water potential at turgor loss (Ψ_l,tlp) and the osmotic potential at full turgidity (Ψ_π,sat) were higher for the new leaves than for the surviving leaves, indicating a lower ability to maintain leaf cell turgor against leaf dehydration in the new leaves. Of the 100 shoots, 13 died after the emergence of new leaves. Stepwise logistic regression revealed that the probability that the long shoots would die generally increased with the emergence of new leaves, with increasing shoot height. This result suggests that the combined effect of the vulnerability of newly emerged leaves and low water availability, associated with higher shoot positions within the crown, caused shoot mortality. Based on our results, some possible mechanisms for mortality in severely defoliated B. maximowicziana are discussed.     

Population structure and dynamics of the clonal dwarf-shrub Linnaea borealis

Abstract. The demography of the long-lived clonal dwarf-shrub Linnaea borealis was studied during four years in a coniferous forest in central Sweden. The main object was to infer patterns of temporal variation in population dynamics of this species. The shoot population is organized in fragments, i.e. physically connected systems of shoots partly covered by the moss carpet. The age and size structure of the fragment population is described, but shoots are more convenient units for a study of population dynamics. A stochastic model of shoot population dynamics was constructed, and simulations indicated a considerable temporal variation in population size and flowering. Hence, variability as such is an essential aspect of the dynamics of established populations of Linnaea. Simulations of extinction risks revealed that small-sized shoot populations (ca. 250 shoots) are likely to be long-lived when experiencing environmentally induced demographic variation of the range observed. Mortality agents for established genets, such as large-scale disturbances, were not incorporated in the models. Some implications of variable population growth rates in clonal plants in woodlands are discussed.     

Seasonal dynamics of shoot biomass of dominant clonal herb species in an oak–hornbeam forest herb layer

Seasonal fluctuations of light availability, nutrient concentrations, and moisture affect plant population traits like density, standing biomass, and flowering. We analyzed seasonal changes of density and shoot biomass of the four most frequent herb species growing in an oak–hornbeam forest community, i.e., Anemone nemorosa, Ficaria verna, Galeobdolon luteum, and Galium odoratum. In 2010 and 2011 plant biomass was harvested from 7 to 10 randomly situated square sample plots (0.36 m²) in the homogenous oak–hornbeam forest community every week in the spring and every two weeks in the summer and autumn. The highest abundance of Anemone nemorosa reached over 1000 shoots per m², of Ficaria verna 459.5 shoots per m², of Galium odoratum 83.6 shoots per m², and of Galeobdolon luteum 98.4 shoots per m² (means for 2010 and 2011, based on all sample plots). We did not observe negative correlation between density and shoot biomass. Growth rates of vegetative shoot biomass amounted to 0.857 mg day^?1 for Anemone nemorosa, 0.467 mg day^?1 for Ficaria verna, 0.722 mg day^?1 for Galium odoratum, and 0.448 mg day^?1 for Galeobdolon luteum (means for 2010 and 2011). Spring ephemerals had much higher densities of shoots than summer-greens. Summer-greens reached higher biomass of individual shoots than spring ephemerals. Flowering shoots constituted only 4, 2, and 11% of all shoots for A. nemorosa, F. verna, and G. odoratum, respectively. More resource availability resulting in high shoot biomass did not translate to higher share of flowering shoots.

     

Translocation of Photoassimilates Between Sister Ramets in Two Rhizomatous Forest Herbs

Autoradiography and liquid scintillation techniques were usedto trace the pattern of photoassimilate translocation in twoperennial forest herbs, Aster acuminatus Michx. and Clintoniaborealis (Ait.) Raf. Vegetative shoots of C. borealis emergeearly each spring from the growing apices of a long-lived rhizomesystem. Vegetative shoots of A. acuminatus emerge in late springfrom rhizomes that decay within 2 years. In both species rametssurvive for only one growing season. Any connected plants aresisters. Mature leaves of these two species were exposed for1 h periods to ¹⁴CO₂ during spring and summer. Radioactivitycould subsequently be detected in exposed leaves (16–84per cent, depending on the season), adjacent above ground plantparts (0.7–23 per cent), roots and rhizomes (4.9–84per cent) and, when present, flowers (4–8.5 per cent).Old rhizomes of C. borealis are prominent storage sites forphotosynthate. In A. acuminatus, no significant translocationbetween sister ramets (i.e. above ground shoots connected bya common rhizome) was observed. In C. borealis, there was small,but consistent translocation between sister ramets (0.2–4per cent). Disturbance of unexposed sister ramets by defoliation,shading or herbivory increased the flow of photoassimilatesto disturbed parts in C. borealis, but not in A. acuminatus.Based on the absence of translocation flow, ramets of A. acuminatusmay be regarded as physiologically independent. Connected rametsof C. borealis show physiological integration. These resultsare correlated with ecological differences between the two species. Aster acuminatus, Clintonia borealis, translocation, ramet, vegetative reproduction, forest herb     

Does morphological plasticity of the <Emphasis Type="Italic">Phalaris arundinacea</Emphasis> canopy increase invasiveness?

Morphological plasticity could facilitate invasions of wetland plants into areas that experience increased durations of flooding and eutrophication. We explored canopy plasticity of Phalaris arundinacea, an aggressive invader of wetlands, as it differentially invaded wet prairie mesocosms under 3 flooding durations and 3 levels of nutrient addition. Phalaris grew as a sward with intermittent and early-season flooding but shifted to tussocks under constant flooding. These two growth forms differed by >20% in several canopy ratios. Clones that formed tussocks produced 45% more shoots per unit biomass (P = 0.007) and a 25% higher ratio of total shoot length to biomass (P = 0.04). Lighter-weight shoots supported 33% fewer leaves and, consequently, had 35% less leaf area per shoot height (P < 0.002). Tussocks developed a continuous mat of adventitious roots, with root mats reaching 20.9 ± 0.6 cm in diameter and 4.7 ± 0.3 cm in height over two growing seasons. While forming tussocks, Phalaris tolerated longer durations of flooding and more than doubled its aboveground biomass. Invasions occurred rapidly, with Phalaris exceeding 75% canopy cover and accounting for 66% of the total aboveground biomass under constant flooding. Early-season flooding increased the lateral spread of individual shoots. High nutrient addition produced shoots that were 27% taller and 50% heavier (P < 0.02), with 81% more leaf area (P < 0.0003) than shoots that received no nutrients. Consequently, under early-season flooding with high nutrient additions, Phalaris was primed to invade, nearly doubling its proportion of the total aboveground biomass and exceeding 50% canopy cover during year two.     

Effects of salinity and cutting on the development of Phragmites australis

The effects of increased salinity and cutting the above ground biomass on the growth of Phragmites australis were evaluated by investigating four experimental reed stands grown in outdoor tanks. Two stands were treated with 30 salinity and the other two stands with freshwater; one stand of each treatment was cut to 20 cm during the second growing season. Growth conditions were observed until all the plants were dead at the end of the second year. The number of shoots emerged from the freshwater-treated stand was about 70% higher than that of the saltwater-treated stand. The number of shoots emerged from cut plant stands were markedly lower than uncut stands. The average shoot height was negatively affected by salinity and shoots that emerged after cutting further decreased in height. The average number of leaves on a shoot was not significantly affected by salinity, but reduced by cutting in both treatments. Leaf length, width and the distance between leaves were decreased by both salinity and cutting. In the freshwater-treated uncut stand more than 50% of the shoots formed panicles, but this proportion was reduced to 6% by salinity, to 15% by cutting, and to 0% by the combination of salinity and cutting. This study showed again that salinity reduces the growth of aboveground components. The growth, however, was most severely retarded by cutting combined with salinity, which has many implications for better management of P. australis stands.