For she that hath, to her shall be given…Implications of flowering in Anemone nemorosa L

We looked for life‐history trade‐offs between flowering, vegetative growth and somatic maintenance in the common woodland herb Anemone nemorosa. A. nemorosa forms a horizontal rhizome system consisting of previously formed annual segments and terminated by a flowering or non‐flowering shoot. Resources acquired by the aboveground parts are used for flowering, seed production, storage and growth of the annual segments. Resources stored in the rhizome during the growing period are used for preformation of buds, somatic maintenance between two growing periods and development of aboveground parts in the following spring. We hypothesised that the decision to invest in flower buds depends on the amount of resources stored in the recently formed annual segment. We also hypothesised a trade‐off between flowering and segment growth and, finally, as a consequence, we expected individual rhizomes to alternate between the flowering and the non‐flowering state. We found that segments producing flower buds were significantly longer than non‐flowering segments, indicating that resource level influences the function of the preformed buds. Contrary to our expectations, we found flowering rhizomes produced longer annual segments than non‐flowering rhizomes. We suggest the larger leaf area of flowering rhizomes and occasional abortion of flowers or seeds as possible mechanisms behind this pattern. Our study shows that even though the decision to produce a flower bud is taken in another time‐frame than that in which the actual flowering and fruiting takes place, an ostensibly inexpedient decision is changed to a neutral or even an advantageous incident.     

Nutrient enrichment enhances hidden differences in phenotype to drive a cryptic plant invasion

Many mechanisms of invasive species success have been elucidated, but those driving cryptic invasions of non‐native genotypes remain least understood. In one of the most successful cryptic plant invasions in North America, we investigate the mechanisms underlying the displacement of native Phragmites australis by its Eurasian counterpart. Since invasive Phragmites’ populations have been especially prolific along eutrophic shorelines, we conducted a two‐year field experiment involving native and invasive genotypes that manipulated nutrient level and competitor identity (inter‐ and intra‐genotypic competition) to assess their relative importance in driving the loss of native Phragmites. Inter‐genotypic competition suppressed aboveground biomass of both native and invasive plants regardless of nutrient treatment (～ 27%), while nutrient addition disproportionately enhanced the aboveground biomass (by 67%) and lateral expansion (by > 3 × farther) of invasive Phragmites. Excavation of experimental plots indicated that nutrient addition generates these differences in aboveground growth by differentially affecting rhizome production in invasive vs native plants; invasive rhizome biomass and rhizome length increased by 595% and 32% with nutrient addition, respectively, while natives increased by only 278% and 15%. Regardless of nutrient level, native rhizomes produced twice as many roots compared to invasives, which field surveys revealed are heavily infected with mycorrhizal symbionts. These results suggest that native Phragmites competes well under nutrient‐limited conditions because its rhizomes are laden with nutrient‐harvesting roots and mycorrhizae. Invasive Phragmites’ vigorous aboveground response to nutrients and scarcity of lateral roots, in contrast, may reflect its historic distribution in eutrophic Eurasian wetlands and correspond to its prevalence in New England marshes characterized by elevated nutrient availability and relaxed nutrient competition. These findings reveal that discrete differences in phenotype can interact with anthropogenic modification of environmental conditions to help explain the success of cryptic invaders.     

Do rhizome severing and shoot defoliation affect clonal growth of Leymus chinensis at ramet population level?

Leymus chinensis (Trin.) Tzvel. is a perennial species of Gramineae, usually subject to defoliation from grazing and mowing. We examined whether shoot defoliation and rhizome severing affected rhizome and ramet growth, and vegetative bud outgrowth of L. chinensis ramet populations. We also tested the hypothesis that clonal growth of the ramets subject to defoliation would benefit from clonal integration between interconnected ramets besides from possible compensatory growth. To 48 experimental plots, we applied six treatments resulting from interactions between two rhizome connection states (unsevered/severed) and three defoliation regimes (non-defoliated, mildly-defoliated and heavily-defoliated). Defoliation affected rhizome growth and bud outgrowth, but had little effect on shoot growth. Mild and heavy defoliation exerted similar effects on rhizome growth. Only heavy defoliation significantly reduced bud outgrowth while mild defoliation did not. The fact that shoot growth did not change after defoliation and that the bud numbers remained unchanged after mild defoliation suggest that the compensatory response enable the species to tolerate grazing to some extent. Neither rhizome severing nor the interaction of rhizome severing and defoliation had effect on any tested variables. Lack of the effect of rhizome severing falsified the first half of our hypothesis, that is, clonal integration was unimportant in our experiment. The probable reasons were suspected to be the short duration of the experiment and/or the buffer effect of carbohydrate reserves in rhizomes for shoot growth and bud production in time of defoliation.     

Mathematical inference of the underground clonal growth of Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae)

Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae) can spread by sexual or vegetative propagation. The choice of strategy likely depends on the environmental conditions. The rhizome is the organ of vegetative reproduction; hence, it is crucial to understand its development. Unfortunately, it is hardly possible to investigate rhizome morphology directly, since E. helleborine is a protected species in most European countries. The goal of our investigation was to infer the growth patterns of underground parts of an orchid population from long-term annual observations of its aboveground shoots. We implemented the Minimum Spanning Tree method to determine a likely set of underground connections between shoots and to simulate the annual growth of new rhizomes. Furthermore, we modelled the spatial distribution of shoots with a density kernel estimator to compare the density gradients with the direction of growth of the rhizomes. Observed shoot numbers fluctuated between 72 and 183 from year to year. Our results suggest that (1) vegetative reproduction prevails in the studied population, (2) the population consists of about a dozen clones with a diameter of up to 6 m, (3) rhizomes produce up to five new shoots at one branch end per year, (4) rhizomes develop in the direction of decreasing population density, and (5) nodes of rhizomes may produce new offshoots after up to 7 years of dormancy.     

Dynamics of growth,carbon and nutrient translocation in Zizania latifolia

We studied the seasonal resource dynamics between organs of wild rice (Zizania latifolia (Griseb.) Turcz. ex Stapf.) to obtain a better understanding of its growth dynamics, carbon and nutrient translocation. The results of observation from January 2002 to February 2004 showed the shoot density markedly increased after emergence of shoots at the end of March until May (up to 800 ind/m²). However the shoot mortality due to self-thinning reduced the total new shoots by more than 70% by the end of July. Thereafter, the shoot density was nearly constant with the aboveground biomass peaking at the end of August. In the late winter, the rhizome biomass declined by respiration loss to about 25% of its peak value. Meanwhile the decline in rhizome reserves from January to the end of April was about 20%. This small reduction compared with other perennial emergent species implies that there is a lower contribution of rhizome reserves to support new shoot formation. The initial heterotrophic growth of new shoots based on the rhizome resources lasted for a short period, then switched to autotrophic growth at the end of April or the beginning of May. Thus, in most periods of foliage development, nutrients were obtained mostly from soil through uptake by roots, not through resource allocation of the rhizome. In autumn, the standing dead shoots retained most of the nutrients and carbohydrates without translocating downwards. This suggests that in practice, the plant can remove nutrients from sediment more efficiently than other emergent plants.     

An individual‐based model of seed‐ and rhizome‐propagated perennial plant species and sustainable management of Sorghum halepense in soybean production systems in Argentina

Perennial plants which propagate through both seeds and rhizomes are common in agricultural and nonagricultural systems. Due to their multifaceted life cycle, few population models are available for studying such species. We constructed a novel individual‐based model to examine the effects of ecological, evolutionary, and anthropogenic factors on the population dynamics of perennial species. To exemplify the application of the model, we presented a case study of an important weed, Sorghum halepense (L.) Pers. (Johnsongrass), in soybean productions in Argentina. The model encompasses a full perennial weed life cycle with both sexual (seeds) and asexual (rhizomes) propagations. The evolution of herbicide resistance was modeled based on either single genes or quantitative effects. Field experiments were conducted in the species' native environment in Argentina to parameterize the model. Simulation results showed that resistance conferred by single‐gene mutations was predominantly affected by the initial frequency of resistance alleles and the associated fitness cost. Population dynamics were influenced by evolved resistance, soil tillage, and rhizome fecundity. Despite the pivotal role of rhizomes in driving the population dynamics of Johnsongrass, most herbicides target the aboveground biomass, and chemical solutions to control rhizomes are still very limited. To maintain effective (short‐term) and sustainable (long‐term) weed management, it is recommended to combine soil tillage with herbicide applications for suppressing the rhizomes and delaying the evolution of resistance. This novel model of seed‐ and rhizome‐propagated plants will also be a useful tool for studying the evolutionary processes of other perennial weeds, cash crops, and invasive species.     

Dynamics of Vaccinium myrtillus patches in mountain spruce forest

Abstract. The dynamics of Vaccinium myrtillus (bilberry) patches were studied in spruce (Picea abies) forest at the higher montane and lower subalpine level in the Tarentaise valley (Savoy, France). Although aerial parts of the shrub may give some indication of age and density of the patches, the annual growth and death of rhizomes are better indicators of patch dynamics. In some cases, dead and young rhizomes may occur simultaneously in the same patch, indicating that growth and death proceed continuously on the inside of dense patches. At mountain sites, slope position of plants significantly influences rhizome growth. A theoretical model of growth of bilberry patches is proposed which takes into account the anisotropic influenceof slope and the postulated independence of different rhizome units belonging to the same individual.     

Age-specific seasonal storage dynamics of<Emphasis Type="Italic">Phragmites australis</Emphasis> rhizomes: a preliminary study

Age-specific seasonal rhizome storage dynamics of a wetland stand of Phragmites australis (Cav.) Trin. ex Steud. in Japan, were investigated from April to October 2000. For each sampling date, above- and below-ground biomass and age-specific rhizome bulk density, ?_rhiz were measured. Seven rhizome age classes were recognized, from <1 year to six years old, based on their position within the branching hierarchy as main criteria and rhizome color, condition of nodal sheaths and condition of the shoots attached to vertical rhizomes as secondary criteria. P. australis stand was moderately productive, having a net aerial and below-ground production of 1980 and 1240 g m^?2, respectively, and a maximum mean shoot height of 2.33 ± 0.12 m. In spring, shoot growth started at the expense of rhizome reserves, decreasing the rhizome biomass as well as ?_rhiz. Both parameters reached the seasonal minimum in May followed by a subsequent increase, indicating a translocation of reserves to rhizomes from shoots after they become self supporting. For each sampling date, ?_rhiz increased with rhizome age. Given that the quantity of reserves remobilized by the rhizomes for spring shoot growth, as assessed by the drop in bulk density from April to May, were positively correlated (r = 0.97, P < 0.05) with rhizome age, it is proposed that for spring shoot formation older rhizomes remobilize stored reserves more actively than younger ones. Given that the accumulation of rhizome reserves (rise in bulk density) from May to August, May to September or May to November was negatively correlated (r = 0.97, 0.92 and 0.87, respectively, P < 0.05) with rhizome age, it seemed possible that younger rhizomes were ‘recharged’ at a higher rate than older ones. These resource allocation mechanisms pertaining seasonal rhizome storage dynamics are of paramount importance in formulating management and conservation strategies of wetlands and aquatic habitats. Our results indicate that a harvest of above-ground biomass from May to June would be more effective in reducing the growth than a harvest in July to August or later, when rhizome reserves have already been replenished. However, the latter may remove a larger shoot bound nutrient stock, still preserving a healthy stand for the subsequent years.     

Rhizome development ofPhragmites australis in a reed community

Variations in the height, shoot density, biomasses of above- and below-ground parts and rhizome distributions ofPhragmites australis were investigated along a line-transect in a reed community at Yufutsu Mire, Hokkaido. Relationships of performance of the reed plants to soil conditions and species compositions were also examined. Three types of rhizome development were recognized in reed plants; (1) the central part of the reed community, characterized by well developed rhizomes and dense aerial shoots, (2) the intermediate part, characterized by development of rhizomes along both the peat and surface layers and very dense aerial shoots, and (3) the marginal part, characterized by development of rhizomes only along the peat layer and sparse aerial shoots. Observation showed that rhizomes in the surface layer actively produced aerial shoots, whereas rhizomes in the peat layer contributed to the spreading of their distribution range. With the growth of rhizomes, organic debris originating from dead rhizomes and roots accumulated in the mineral soil to promote organic soil formation. In dense parts of the reed stand, species composition was poor because of the shading and litter accumulation by reed plants. The effects of microtopography and water level on the establishment of reed seedlings were also considered.     

Seasonal patterns of partitioning and remobilization of 14C in the invasive rhizomatous perennial Japanese knotweed (Fallopia japonica (Houtt.) Ronse Decraene)

Resource partitioning between shoot growth, storage and reproduction is poorly understood in many clonal plant species. This study documents seasonal patterns of growth, ¹⁴C-labelled photoassimilate distribution and remobilization in the invasive rhizomatous species Fallopia japonica (Japanese knotweed). Biomass accumulation above- and below-ground in F. japonica was rapid. By September, rhizome biomass had increased 18-fold from the initial harvest in May (representing 48% of total plant biomass) and this was maintained over winter. Patterns of ¹⁴C allocation from F. japonica shoots labelled at different times of year show that as the season progressed, the rhizomes became an increasingly important sink for current assimilate (the percentage of ¹⁴C recovered from rhizomes was 35% in August and 67% in September) and the corresponding retention of assimilate by established shoots declined. The percentage of ¹⁴C exported to roots was greatest in August. Relatively little photoassimilate was exported to other shoots on the plant, or to flowers. Recycling of photoassimilate was fairly tight in this species and ¹⁴C fixed by shoots in early May 1999 or September 1999 was remobilized to the rhizome prior to shoot senescence and death. Some of this ¹⁴C was then remobilized to new shoots early the following spring. These characteristics may contribute to the success of F. japonica in colonizing a variety of contrasting habitats, often with serious management implications.     

Elevated atmospheric CO2 stimulates aboveground biomass in a fire-regenerated scrub-oak ecosystem

The effect of elevated atmospheric CO₂ concentration (C_a) on the aboveground biomass of three oak species, Quercus myrtifolia, Q. geminata, and Q. chapmanii, was estimated nondestructively using allometric relationships between stem diameter and aboveground biomass after four years of experimental treatment in a naturally fire‐regenerated scrub‐oak ecosystem. After burning a stand of scrub‐oak vegetation, re‐growing plants were exposed to either current ambient (379 µL L^?1 CO₂) or elevated (704 µL L^?1 CO₂) C_a in 16 open‐top chambers over a four‐year period, and measurements of stem diameter were carried out annually on all oak shoots within each chamber. Elevated C_a significantly increased aboveground biomass, expressed either per unit ground area or per shoot; elevated C_a had no effect on shoot density. The relative effect of elevated C_a on aboveground biomass increased each year of the study from 44% (May 96–Jan 97), to 55% (Jan 97–Jan 98), 66% (Jan 98–Jan 99), and 75% (Jan 99–Jan 00). The effect of elevated C_a was species specific: elevated C_a significantly increased aboveground biomass of the dominant species, Q. myrtifolia, and tended to increase aboveground biomass of Q. chapmanii, but had no effect on aboveground biomass of the subdominant, Q. geminata. These results show that rising atmospheric CO₂ has the potential to stimulate aboveground biomass production in ecosystems dominated by woody species, and that species‐specific growth responses could, in the long term, alter the composition of the scrub‐oak community.     

Effects of lead contamination on the clonal propagative ability of Phragmites australis (common reed) grown in wet and dry environments

Clonal propagation is important for the survival and maintenance of the common reed Phragmites australis. Pot culture experiments were conducted to investigate the effects of lead (Pb) concentration (0, 500, 1500, 3000, 4500 mg·kg^?1) and water stress on the clonal reproductive ability of this species. The Pb concentration found in plant organs, in decreasing order, was roots >shoots >rhizomes. There was a negative relationship between the growth of clonal propagative modules (excluding axillary shoot buds) and Pb concentrations, which caused a decrease in biomass, rhizome growth and number of axillary and apical rhizome buds. Daughter axillary shoots exhibited a tolerance strategy, with no significant change in their number; the axillary and apical rhizome buds, daughter apical rhizome shoots and rhizomes exhibited compensatory growth during the late stage of Pb (excluding 4500 mg·kg^?1) treatment in a wet environment. Pb applications above 500 mg·kg^?1 reduced these parameters significantly in the drought treatment, except for the number of axillary shoot buds, which did not change. Our results indicate that clonal propagative resistance to Pb contamination can occur via tolerance strategies, compensatory growth and a Pb allocation strategy, enabling these reeds to maintain population stability in wet environments. However, clonal modular growth and reproductive ability were inhibited significantly by the interaction between drought and Pb, which would cause a decline in P. australis populations in a dry environment. Lead concentrations of 4500 and 500 mg·kg^?1 in soils might meet or exceed the Pb tolerance threshold of clonally propagated reeds in wet and dry environments, respectively.     

Growth forms of Leymus chinesis (Poaceae) at the different developmental stages of the natural population

The manner in which the density of Leymus chinensis increases from a single plant to a dominant population can be understood by tracing the development of a population from early to late stages. Parent shoot density, above‐ground dry weight, spike density, heading rate and spike dry weight, density of spreading shoots (buds/daughter shoots in apical/axillary rhizomes) and clumping shoots (buds/daughter shoots in axillary parent shoots), and young rhizome length and weight were investigated in the same quadrats for a low density/early stage (LE) population and a high density/late stage (HL) population. Clonal growth (buds/daughter shoots formation) and sexual reproduction (spikes formation) increased while rhizome storage (young rhizome weight) decreased during the transition from LE to HL. In a LE population an outward occupation strategy was employed, with a high proportion of spreading shoots. As the population density gradually increased until HL, an inward consolidation strategy increasing shoot amount in previously occupied areas, was adopted. This was characterized by a high proportion of clumping shoots. Interestingly, the trade‐off between spreading and clumping shoots can be adjusted by the duration of young rhizome elongation during a growth season. In other words, compared with a HL population, a LE population shortened the duration of young rhizome elongation during the growth season, which resulted in more time for the production of axillary spreading shoots along the rhizomes, and high amounts and proportions of total spreading shoots. The special growth patterns, that is, trade‐offs among growth forms, allow L. chinensis to establish dominant populations throughout the eastern Eurasian Steppe.     

The effects of 1-naphthaleneacetic acid and N6-benzyladenine on the growth of Cymbidium forrestii rhizomes in vitro

An Asiatic orchid, Cymbidium forrestii, was clonally propagated using seed-derived rhizomes as explants. The rhizomes were cultured and proliferated on Murashige and Skoog medium supplemented with various growth substances. Auxins stimulated rhizome growth by increasing branching and fresh weight of the explant, with 1-naphthaleneacetic acid (NAA) being the most effective auxin. All auxins tested suppressed normal shoot formation. The apical meristem of the rhizome reacted to exogenously applied auxin by reducing the cytoplasmic zone of the apical meristem and causing meristem derivatives to rapidly differentiate into vacuolated parenchyma cells. Leaf formation and development was retarded in the presence of auxin. Cytokinins generally reduced rhizome growth and the number of branches, but benzyladenine (BA) can induce shoot formation in vitro. BA induced the cytoplasmic zone of the apical meristem to enlarge and enhanced leaf development. A 5% (w/v) sucrose concentration was most effective in shoot induction when combined with 5 mg1^-1 BA. Activated charcoal promoted rhizome growth; however, shoot formation was inhibited.     

DEVELOPMENTAL STUDIES ON THE COENOCYTIC ALGA, CAULERPA SERTULARIOIDES

Thallus growth and development in the coenocyti alga Caulerpa sertularioides (Gmelin) Howe have been studied quantitatively. In unsupplemented seawater at 26 C and in a 12:12 hr light/dark cycle, new rhizomes formed near the old growing points on thalli transplanted from the ocean. Adjacent to the apices of new rhizomes, rhizoids were produced downward, regularly spaced and at a rate of about 1.5/day. Upright. shoots developed irregularly in acropetal succession behind the tips of either the parent or the new rhizomes, which arose as branches of the old. Rates of rhizome, rhizoid, and upright shoot elongation were 0.4, 0.81, and 0.54 cm/day, respectively. Thalli survived up to 2 months in unsupplemented seawater. Long-term growth was obtained by varying culture conditions. A substratum of sand, apparently rich in microorganisms, produced long-term thallus growth in seawater, and the form of development changed so that upright shoot formation was promoted and rhizome elongation halved. Similar effects were elicited by indole-3-acetic acid, 5 × 10^-5, M in seawater and by sap expressed from C. racemosa or C. sertularioides and added to seawater at 2.5–10% by volume. The regulation of development in an algal coenocyte is discussed and analogies with regulation in multicellular plants are drawn.     

Variable responses of native eelgrass Zostera marina to a non-indigenous bivalve Musculista senhousia

The transport and establishment of non-indigenous species in coastal marine environments are increasing worldwide, yet few studies have experimentally addressed the interactions between potentially dominant non-native species and native organisms. We studied the effects of the introduced mussel Musculista senhousia on leaf and rhizome growth and shoot density of eelgrass Zostera marina in San Diego Bay, California. We added M. senhousia over a natural range in biomass (0–1200 g dry mass/m²) to eelgrass in transplanted and established beds. The effects of the non-indigenous mussel varied from facilitation to interference depending on time, the abundance of M. senhousia, and the response variable considered. Consistent results were that mussel additions linearly inhibited eelgrass rhizome elongation rates. With 800 g dry mass/m² of M. senhousia, eelgrass rhizomes grew 40% less than controls in two eelgrass transplantations and in one established eelgrass bed. These results indicate that M. senhousia, could both impair the success of transplantations of eelgrass, which spread vegetatively by rhizomes, and the spread of established Z. marina beds to areas inhabited by M. senhousia. Although effects on leaf growth were not always significant, in August in both eelgrass transplantations and established meadows leaf growth was fertilized by mussels, and showed a saturation-type relationship to sediment ammonium concentrations. Ammonium concentrations and sediment organic content were linear functions of mussel biomass. We found only small, non-consistent effects of M. senhousia on shoot density of eelgrass over 6-month periods. In established eelgrass beds, but not in transplanted eelgrass patches (≈0.8 m in diameter), added mussels suffered large declines. Hence, eelgrass is likely to be affected by M. senhousia primarily where Z. marina beds are patchy and sparse. Our study has management and conservation implications for eelgrass because many beds are already seriously degraded and limited in southern California where the mussel is very abundant. Received: 31 May 1997 / Accepted: 4 September 1997     

Central Die-back of Monoclonal Stands of Reynoutria japonica in an Early Stage of Primary Succession on Mount Fuji

Reynoutria japonica is a common perennial pioneer species onJapanese volcanoes. In a volcanic desert (1500m above sea level)on Mount Fuji (3776m), central Japan, this species forms circularstands (patches). As a patch develops, shoot density decreasesin its centre (‘central die-back’). The centraldie-back has been considered a key process in the early stagesof primary succession, though its mechanism has been unknown. The pattern of patch development, population dynamics of aerialshoots, and growth patterns of below-ground organs were analysedin order to investigate the mechanism of die-back, and the followingtraits are clarified: (1) central die-back areas occur in mostsmall patches (approx. 1m²) without later successional species;(2) shoot characteristics are dependent both on their positionwithin a patch and on patch size; (3) despite the large differencesin shoot density, neither time course of shoot growth nor theirmortality differs between the centre and periphery of patches;and (4) rhizomes ofR. japonicagrow outwards with regular sympodialbranching. From these results, it is concluded that neither interspecificnor intraspecific competition is likely to be a primary causeof the die-back phenomenon, but that central die-back is broughtabout intrinsically by the growth pattern of the rhizome systems.We also discuss the importance of the central die-back in facilitatingestablishment of later successional species in the early stagesof primary succession. Clonal plant; central die-back; competition; facilitation; Japanese knotweed; Mount Fuji; primary succession;Reynoutria japonica ; rhizome growth; volcanic desert     

Demography of Carex brevicuspis (Cyperaceae) rhizome populations: a wetland sedge that produces both elongated and shortened rhizomes

The wetland sedge Carex brevicuspis reproduces vegetatively by producing short rhizomes to form clumping ramets phalanx) and long rhizomes to form spreading ramets (guerrilla), resulting in a combined growth form. As an initial step towards understanding the adaptation of Carex growth strategies to seasonal fluctuations in wetland habitats, we investigated the density and composition of C. brevicuspis rhizome populations immediately after flooding (November), in winter (January), in spring (March), and before flooding (May) in the Dongting Lake wetlands, China. The total rhizome density peaked in winter and was lowest before flooding. A large rhizome population in winter may enable C. brevicuspis to survive the seasonal cold weather and recruit a shoot population in the spring. A small rhizome population before flooding may optimize reproductive allocations and be a strategy for enduring the long flooding season. Regardless of date, short rhizomes comprised the majority of the rhizome population (73.0% in March to 98.2% in May). This indicates that C. brevicuspis primarily uses a phalanx growth strategy to utilize locally abundant resources in wetlands. The percentage of long rhizomes in the rhizome population varies significantly between seasons (1.8% in May to 27.0% in March), indicating that growth form also changes with seasonal fluctuation of wetland habitats. The results show that C. brevicuspis may adapt to seasonal changes in wetland habitats through changes in demography of rhizome populations.     

Factors Affecting Revegetation of Carex lacustris and Carex stricta from Rhizomes

The revegetation of sedge meadows has been problematic because natural recolonization does not occur under many circumstances and because planted propagules often fail to reestablish successfully. In this study, detached rhizomes of Carex lacustris Willd. and Carex stricta Lam. were transplanted in both fall (September) and spring (May) into three experimental wetlands to determine the effects of both planting season and hydrology on survival and establishment. Each experimental wetland had the same mean water depth across 5% slopes, but one had a constant water depth (0.5 m) throughout the growing season, another fell from a mean depth of 0.75 m to 0.25 m, and a third rose from a mean depth of 0.25 m to 0.75 m. Initial rhizome survival, shoot growth, and soil characteristics were recorded over 2 years. Neither planting proved successful (6.9% versus 0.5%) for C. stricta, a tussock-forming sedge. For C. lacustris, a sedge with spreading rhizomes, spring planting had greater rhizome survival (53.2% survival) than fall planting (0.7%). Since both species initiate new shoots in the fall, they are susceptible to transplant failure during this season. The highest survival rates (71–100%) and plant production (736.0 and 494.5 g/m²) for C. lacustris occurred near the water’s edge in both the constant and falling basins. In the rising basin, establishment and growth of this species was high at all water depths (71–96%; 399 g/m²). C. lacustris grew optimally at the same elevations where rhizome survival was greatest, suggesting that shoots are more sensitive to early-season than late-season water levels.     

Contrasting seasonal overlaps between primary and secondary growth are linked to wood anatomy in Mediterranean sub‐shrubs

Whole‐plant approaches allow quantification of the temporal overlap between primary and secondary growth. If the amount of time available to grow is short, there may be a high temporal overlap between shoot growth and wood formation. We hypothesise that such overlap depends on the duration of the growing season and relates to wood anatomy. We evaluated wood anatomy, shoot longitudinal and radial growth rates, fine root production and the concentrations of non‐structural carbohydrates (NSC) in the wood of six sub‐shrub species growing in sites with contrasting climatic conditions (Lepidium subulatum, Linum suffruticosum, Salvia lavandulifolia, Satureja montana, Ononis fruticosa, Echinospartum horridum). Sub‐shrub species living in sites with a short growing season displayed a high overlap between aboveground primary and secondary growth and formed wide vessels, whereas species from the warmest and driest sites presented the reverse characteristics. The highest overlap was linked to a rapid shoot extension and thickening through the enhanced hydraulic conductivity provided by wide vessels. The reductions in NSC concentrations when growth peaked were low or moderate, indicating that sub‐shrubs accumulate NSC in excess, as do trees. The temporal overlap among primary and secondary growth in woody plants may be connected to the duration and rates of shoot and wood growth, which in turn depend on the vessel lumen area.