Constraints on Sedge Meadow Self‐Restoration in Urban Wetlands

Invasive plants and urban run‐off constrain efforts to restore sedge meadow wetlands. We asked if native graminoids can self‐restore following the removal of Typha × glauca (hybrid cattail), and if not, what limits their recovery? After we harvested Typha and depleted its rhizome starch reserves, Carex spp. expanded vegetatively (approximately 1 m over 2 years) but not by recruiting seedlings. A seedling emergence experiment showed that seed banks were depleted where Typha had eliminated the sedge meadow over a decade ago (based on aerial photo analysis). Carex seedling emergence was 75–90% lower where Carex was absent than where it remained in the plant community, and at least 17 species that were abundant 30 years ago were absent from the seed bank and extant vegetation. By varying hydroperiod, we showed that prolonged flooding prevented emergence of Carex seedlings and that a fluctuating hydroperiod reduced emergence and ultimately killed all Carex seedlings. In contrast, Typha seedlings emerged and survived regardless of hydroperiod. Thus, slow vegetative expansion by Carex, depauperate seed banks, and altered hydroperiods all constrain self‐restoration. To compensate for multiple constraints on self‐restoration, we recommend a long‐term management approach that capitalizes on flooding and the capacity of Carex spp. to regrow vegetatively. We suggest annually harvesting swaths of Typha at the edges of clones, before or during flood events, to allow gradual, vegetative self‐restoration of Carex spp.     

A Test of Two Annual Cover Crops for Controlling Phalaris arundinacea Invasion in Restored Sedge Meadow Wetlands

Abstract Rapid establishment by aggressive plants such as Phalaris arundinacea (reed canarygrass) often interferes with sedge meadow establishment in restored prairie pothole wetlands in the mid‐continental United States. Introducing a cover crop during community establishment might suppress P. arundinacea invasion in restored prairie potholes by reducing resource availability. We evaluated two potential cover crops, Echinochloa crusgalli (barnyardgrass) and Polygonum lapathifolium (nodding smartweed), for suppressing P. arundinacea invasion in an experimental wetland using replacement series competition experiments. Further, we assessed the effects of E. crusgalli and P. lapathifolium on sedge meadow establishment by sowing Carex hystericina, a common wetland sedge, as a third species at a constant density in the replacement experiments. Echinochloa crusgalli, compared with no cover crop, reduced P. arundinacea biomass by more than 1,000 g/m² (65%) after two growing seasons. Polygonum lapathifolium did not affect P. arundinacea biomass. Dense E. crusgalli canopies in the first year and thick E. crusgalli thatch in the second year substantially reduced light availability for P. arundinacea establishment. Echinochloa crusgalli also reduced C. hystericina biomass by more than 1,800 g/m² (99%) after two growing seasons. Carex hystericina biomass was similar in plots sown with E. crusgalli to P. arundinacea monocultures. Neither E. crusgalli nor P. lapathifolium is likely to improve sedge meadow restoration success. These trends were not sensitive to initial sowing density or elevation above water level. Without methods to suppress P. arundinacea invasions, sedge meadow restorations may often fail. Thorough site preparation to remove P. arundinacea propagule sources before restoration is essential.     

Tools for <Emphasis Type="Italic">Carex</Emphasis> revegetation in freshwater wetlands: understanding dormancy loss and germination temperature requirements

Carex is a globally distributed genus with more than 2000 species worldwide and Carex species are the characteristic vegetation of sedge meadow wetlands. In the mid-continental United States, Carex species are dominant in natural freshwater wetlands yet are slow to recolonize hydrologically restored wetlands. To aid in Carex revegetation efforts, we determined the dormancy breaking and temperature germination requirements of 12 Carex species. Seeds were cold stratified at 5/1°C for 0–6 months and then incubated in light at 5/1°C, 14/1°C, 22/8°C, 27/15°C, or 35/30°C. We found that all Carex species produced conditionally dormant seeds. The optimal temperature for germination for all but three species was 27/15°C. As is the case in other species with physiological dormancy, cold stratification increased germination percentages, broadened the temperature range suitable for germination, and increased germination rates for most species, but the magnitude of the effects varied among species. Many species germinated to 80% at 27/15°C without cold stratification and at 22/8°C with ≤1 month of stratification but required much longer stratification (up to 6 months depending on the species) to germinate to 80% at 14/1°C and 35/30°C. Our findings illustrate how a stratification pretreatment can greatly benefit Carex seed sowing efforts by triggering rapid germination to higher percentages. We recommend that cold stratification be targeted towards species with strong dormancy or used across a wider range of species when seed supplies for restoration are limiting. For Carex revegetation, establishing Carex canopies rapidly may help to prevent the invasion of undesirable species such as Phalaris arundinacea.     

Temperature requirements for dormancy break and seed germination vary greatly among 14 wetland Carex species

We evaluated dormancy loss in seeds of 14 Carex species (C. atherodes, C. brevior, C. comosa, C. cristatella, C. cryptolepis, C. granularis, C. hystericina, C. lacustris, C. pellita, C. scoparia, C. stipata, C. stricta, C. utriculata, C. vulpinoidea) under growing season and stratification conditions and determined the temperature requirements for germination. Seeds were germinated for 1 year at a diel temperature regime (5/1 °C, 14/1 °C, 22/8 °C, or 27/15 °C) or a seasonal regime (seeds moved among the four diel regimes to mimic seasonal temperatures). All species had conditionally dormant seeds at maturity. The optimal temperature for germination of most species was 27/15 °C. The 14 species were grouped by their seed viability, dormancy, and germination with a Seed Regeneration Index (SRI; range 0–1) using the results of this study and a previously published paper on stratification effects on Carex seed dormancy and germination. The eight species that had an SRI value >0.5 (C. brevior, C. comosa, C. cristatella, C. cryptolepis, C. hystericina, C. scoparia, C. stipata, C. vulpinoidea) had high seed viability (>60%) and required little to no stratification to germinate readily over a broad range of temperatures. The six species with an SRI value <0.5 (C. atherodes, C. granularis, C. lacustris, C. pellita, C. stricta, C. utriculata) generally had low seed viability (<50% and often <1%) and required stratification or particular temperatures (35/30 °C or 5/1 °C for C. stricta; 35/30 °C for C. utriculata; 27/15 °C for C. atherodes, C. lacustris, C. pellita; 5/1 °C for C. granularis) for germination ≥50%. These six species will require more attention from restoration practitioners to ensure that there are sufficient viable seeds to meet revegetation goals, that dormancy break is achieved, and that seeds are sown when temperatures are optimal for germination. The different seed germination syndromes that we found for these Carex species likely contribute to variable seed bank formation and emergence patterns, and species coexistence.     

Effect of light on seed germination of eight wetland Carex species

BACKGROUND AND AIMS: In wetland plant communities, species-specific responses to pulses of white light and to red : far-red light ratios can vary widely and influence plant emergence from the seed bank. Carex species are the characteristic plants of sedge meadows of natural prairie wetlands in mid-continental USA but are not returning to restored wetlands. Little is known about how light affects seed germination in these species-information which is necessary to predict seed bank emergence and to develop optimal revegetation practices. The effects of light on germination in eight Carex species from prairie wetlands were investigated. METHODS: Non-dormant seeds of eight Carex species were used to determine the influence of light on germination by examining: (a) the ability of Carex seeds to germinate in the dark; (b) the effect of different lengths of exposures to white light on germination; (c) whether the effect of white light can be replaced by red light; and (d) whether the germination response of Carex seeds to white or red light is photoreversible by far-red light. KEY RESULTS: Seeds of C. brevior and C. stipata germinated >25 % in continuous darkness. Germination responses after exposure to different lengths of white light varied widely across the eight species. Carex brevior required <15 min of white light for > or =50 % germination, while C. hystericina, C. comosa, C. granularis and C. vulpinoidea required > or =8 h. The effect of white light was replaced by red light in all species. The induction of germination after exposure to white or red light was reversed by far-red light in all species, except C. stipata. CONCLUSIONS: The species-specific responses to simulated field light conditions suggest that (a) the light requirements for germination contribute to the formation of persistent seed banks in these species and (b) in revegetation efforts, timing seed sowing to plant community development and avoiding cover crops will improve Carex seed germination.     

Nonequilibrium dynamics of sedge meadows grazed by cattle in southern Wisconsin

Equilibrium theory predicts that after disturbance, ecosystemseventually regain the structural and functional properties characteristic oftheir predisturbance condition. This study tested this idea by examining theeffects of cattle grazing and exclusion on the long-term structuralcharacteristics of sedge meadows in southern Wisconsin. To compare structuralchanges in mean percentage cover and height, repeated measures analysis wasconducted on two sedge meadows over a twenty year period from 1977 to 1997. Onesedge meadow was recovering from cattle grazing (cattle excluded in 1973) andthe other was a reference area (nearly undisturbed). Both of these study siteschanged structurally from 1977 to 1997, supporting non-equilibrium theory.Additional observations were made in a heavily and lightly grazed sedge meadowthat were surveyed in 1977. As based on the positions of subunits in anordination graph produced using Non-Metric Multidimensional Scaling (NMS), therecovery sedge meadow became less structurally similar to the grazed and moresimilar to the reference site over the 20 year study. However, from theperspective of mean maximum height in another NMS analysis, the recovery sedgemeadow became less similar to the reference site over time likely because by1997, a shrub carr of Cornus sericea had developed in therecovery sedge meadow that had been dominated by graminoids and forbs in 1977(mean maximum height: 1977 vs. 1997; 0 vs. 47 cm). Seedlings ofCornus sericea were invading the grazed sedge meadows andin the recovery sedge meadow (cattle excluded 4 years earlier) in 1977. A shrubcarr did not develop in the reference sedge meadow. Changes in the referencesite were relatively minor over this time interval; certain species eitherincreased or decreased in dominance, e.g., Carex strictaincreased in cover (1977 vs. 1997, 20 and 28 mean percentage (%) cover,respectively). A few short-term species of the recovery sedge meadow followedthe tenets of equilibrium theory. These became less common or disappeared4–9 years after cattle exclusion including Asterlanceolatus, Calamagrostis canadensis,Poa compressa, Solidago altissima andVerbena hastata. Some of these species were eaten andlikely spread by the cattle. This study suggests that the progression of sedgemeadow to shrub carr may not be an inevitable outcome of succession but insteadcan be a consequence of past cattle grazing history. Also, because the recoveryand the reference sedge both changed structurally over time, the tenets ofnon-equilibrium theory were supported by this study.     

Factors Affecting Seed Germination and Seedling Establishment of Fen-Meadow Species

Availability of seeds and provision of “safe sites” for seedling recruitment are essential for successful restoration of seminatural grassland communities. Inability to provide species‐specific conditions for seedling recruitment appears to be a major factor limiting establishment of fen‐meadow species on restoration sites. This contention was tested in the field for both germination and establishment conditions for a selection of fen‐meadow species. A Cirsio‐Molinietum fen meadow and an agriculturally semi‐improved species‐poor grass dominated rush pasture were used. Seeds of Carex ovalis, Cirsium dissectum, Molinia caerulea, Succisa pratensis, and Holcus lanatus were sown onto treatments comprising either irrigation or no irrigation, presence or absence of existing vegetation canopy, and presence or absence of soil disturbance. Germination of all except H. lanatus was higher in the fen meadow than in the rush pasture. The fen‐meadow site was less susceptible to drought, provided more light to the seed environment, and showed a stronger day–night variation in relative humidity compared with the rush pasture. All the fen‐meadow species responded strongly to the experimental treatments, whereas H. lanatus showed only a small response. Soil disturbance was the major factor that increased germination. Removal of the vegetation canopy improved germination only in S. pratensis. Conditions affecting survival of seedlings were different from those affecting seed germination. Seedling survival was greater on the fen‐meadow site than on the rush pasture. Canopy presence was the major factor that reduced seedling survival. Few seedlings survived in the presence of the rush pasture canopy. Irrigation and soil disturbance were of minor importance for seedling survival on both sites. Safe sites for seed germination and seedling establishment of fen‐meadow species existed on the fen meadow even without soil disturbance and gap creation. Safe sites for seedling recruitment were not present in the rush pasture. The need for species‐specific definition of safe site characteristics at the two stages of seedling recruitment (i.e., for seed germination and for seedling survival) was demonstrated. The implications of these findings for restoration of seminatural grasslands are discussed.     

Establishment of Carex stricta Lam. seedlings in experimental wetlands with implications for restoration

The loss of Carex dominated meadows due to agricultural drainage in the previously glaciated midcontinent of North America has been extensive. The lack of natural Carex recruitment in wetland restorations and the failures of revegetation attempts underscore the need for information on the establishment requirements of wetland sedges. In this study, seedlings of Carex stricta Lam. were planted in three experimental wetlands in east-central Minnesota, USA to investigate the biotic and abiotic environmental limitations to establishment. Seedlings were planted along an elevational water depth gradient to assess the effects of water depth and water level fluctuation on seedling survival and growth. A different water level fluctuation regime was assigned to each of the experimental wetlands to assess seedling tolerance for seasonal water level changes. The effects of seedling planting density and the presence or absence of non-sedge colonizers on seedling survival and growth were also studied. The experiment was followed for three growing seasons. The results of this study indicate that C. stricta seedlings were sensitive to the timing and duration of inundation during the first growing season. Once established, plants tolerated a broad range of seasonal drying and flooding conditions. Seedling and juvenile growth was slowed by non-sedge colonizers during the first two growing seasons, but by the third growing season, C. stricta was able to out-grow all annual and perennial weeds, except the aggressive perennial, Phalaris arundinacea L. The rapid growth of C. stricta plants, once established, indicates that the use of seedlings is a successful method for (re) introducing this tussock sedge into wetland restorations under a variety of environmental conditions. Comparison with other studies performed under similar conditions suggests that planting of seedlings is a more appropriate method of establishing this species than the use of transplanted rhizomes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Zonation of plant cover and environmental factors in wetlands of the Sanjiang Plain,northeast China

The zonation of depressional and riparian wetlands in the Sanjiang Plain of northeastern China was studied to describe their vegetation composition and environmental variables. We sampled 108 plots in 6 depression and riparian wetlands. Samples were classified into 4 groups using two‐way indicator species analysis (TWINSPAN). Emergent marsh vegetation was characterized by Carex lasiocarpa and C. pseudocuraica, meadow marsh vegetation by tussock species such as Carex appendiculata and C. meyeriana, wet meadow vegetation by Calamangrostis angustifolia and Carex appendiculata, and shrub meadow vegetation by the shrubs Betula fruticosa, Alnus sibirica and Salix rosmarinifolia and the graminids Carex schmidtii and Calamagrostis angustifolia. CCA ordination showed that water table, organic matter and available N were the major factors explaining the vegetation zonation pattern. Compared with other Northern Hemisphere regions, bog and fen vegetation are completely absent due to climatic conditions unfavorable for peat formation. Out of four vegetation types, only the Carex lasiocarpa community and the C. pseudo‐curaica community have been found in other regions. However, at the species level many species are widespread and some species are vicariant or pseudovicariant to other regions in the world. Our study suggests that topography and hydrology may be the most important determinants of the vegetation pattern in this region.     

Achene germination of the spring ephemeroid species Carex physodes in the Gurbantunggut Desert

Much of the seed germination research on Carex has focused on wetland species, and little is known about the species of arid habitats. Here, we investigated seed dormancy and germination of Carex physodes, which is an important component of the plant community of the Gurbantunggut Desert of the Junggar Basin in Xinjiang, China. Our studies included the effects of mechanical and chemical scarification, dry storage, treatment with GA₃, wet‐cold stratification, and burial in the field. No freshly matured achenes germinated over a range of temperature regimes after treatment with GA₃, 6 months of dry storage or removal of part of the endosperm. The mechanical scarification resulted in < 5% achene germination, however, higher percentage of achene germination occurred after removal of the pericarp (60%), H₂SO₄ scarification (30%) or scarification in 10% NaOH (85%). Six and nine months of wet‐cold stratification promoted < 40% achene germination. The optimal germination temperatures ranged from 25/10°C to 35/20°C. Maximum germination after 9 months of burial at a depth of 3 cm in the field was 36%. These results indicate that the seeds have non‐deep physiological dormancy (PD) and that the pericarp contains germination inhibitors and has strong mechanical resistance to germination.     

Identifying Preferential Associates to Initiate Restoration Plantings

Ecological restoration typically aims to re‐establish dominant plant species and their native associates, despite the lack of guidance on which associates to introduce initially. Analysis of naturally occurring plant communities can provide criteria to shorten the list of species that are associated with dominants, in order to focus revegetation efforts on species likely to establish. Using the example of sedge meadows, we evaluated wetland vegetation data from Laurentian Great Lakes wetlands to identify “preferential associates,” that is, species that co‐occur more frequently than expected based on their overall abundance. A total of 176 taxa occurred within the two hundred and thirty‐nine 1 × 1 m² plots in 48 wetlands that contained Carex stricta, a widespread tussock‐forming sedge. Of 58 species that co‐occurred with C. stricta where it was dominant (≥50% plot cover), we identified 26 associates using Bray–Curtis similarities and we determined that 12 of the 26 were preferential using an electivity index. The electivity index identified preferential associates even when they occurred infrequently or had low mean cover per plot. We provide guidance on how to initiate restoration with preferential associates.     

Comparisons of nutrient recovery and specific leaf area variation betweenCarex lasiocarpa var.occultans andCarex thunbergii var.appendiculata with reference to nutrient conditions and shading byPhragmites australis

The distribution of two sedge species was studied in two mires which differ in abiotic environments and in distribution ofPhragmites australis. Carex lasiocarpa var.occultans dominated in nutrient-poor valley mire, andCarex thunbergii var.appendiculata dominated in nutrient-rich flood plain subject to water fluctuations.Phragmites australis grew well in nutrient-rich conditions. The distribution ofC. lasiocarpa showed a strong negative correlation withP. australis coverage, whereasC. thunbergii coverage was not affected byP. australis. The leaf area per dry leaf mass (specific leaf area: SLA) ofC. thunbergii increased with shading byP. australis, but that ofC. lasiocarpa was stable. The SLA flexibility ofC. thunbergii to light interception might enable this species to invadeP. australis patches in nutrient-rich environments. The residual nutrient ratio of nitrogen and phosphorus (the ratio of the residual nutrient content at the end of the growing season to peak nutrient content) in the vegetative ramet ofC. thunbergii was 1.7 times higher than that ofC. lasiocarpa. This low residual ratio may indicate effective nutrient recovery to storage organs. The effective nutrient recovery inC. lasiocarpa might enable this species to grow even in nutrient-poor environments. However, it may be difficult forC. lasiocarpa to expand its habitat to nutrient-rich areas whereP. australis dominates as it is not shade tolerant.     

Restoration Potential of Sedge Meadows in Hand‐Cultivated Soybean Fields in Northeastern China

Sedge meadows can be difficult to restore from farmed fields if key structural dominants are missing from propagule banks. In hand‐cultivated soybean fields in northeastern China, we asked if tussock‐forming Carex and other wetland species were present as seed or asexual propagules. In the Sanjiang Plain, China, we compared the seed banks, vegetative propagules (below‐ground) and standing vegetation of natural and restored sedge meadows, and hand‐cultivated soybean fields in drained and flooded conditions. We found that important wetland species survived cultivation as seeds for some time (e.g. Calamogrostis angustifolia and Potamogeton crispus) and as field weeds (e.g. C. angustifolia and Phragmites australis). Key structural species were missing in these fields, for example, Carex meyeriana. We also observed that sedge meadows restored without planting or seeding lacked tussock‐forming sedges. The structure of the seed bank was related to experimental water regime, and field environments of tussock height, thatch depth, and presence of burning as based on Nonmetric Multidimensional Scaling analysis. To re‐establish the structure imposed by tussock sedges, specific technologies might be developed to encourage the development of tussocks in restored sedge meadows.     

Increased Germination Rates of Baldcypress and Pondcypress Seed Following Treatments Affecting the Seed Coat

The effects of seed-coat removal, warm-water soaking, cold stratification, dry-cold storage and acid treatment on the germination of baldcypress (Taxodium distichum (L.) Rich.) and pondcypress (Taxodium distichum var. nutans (Ait.) Sweet) seed were studied. Removal of the hard seed coat resulted in a prompt and high percent germination, indicating that there was no physiological embryo dormancy. Cold stratification, warm-water soaking and acid treatment did not increase percent germination over the control after a 60-day germination period. They did promote the rate of germination however, having a significantly higher germination at 30 days. Dry-cold storage promoted the initial rate of germination, but resulted in a decreased percent germination below the control after 60 days. The above results suggest that moisture, and not cold, was the crucial component of stratification. The improved germination rate following acid treatment further suggests that any treatment which rendered the seed coat more permeable or softened it was more effective in increasing the rate of germination of baldcypress and pondcypress seed.     

Revegetation of prairie pothole wetlands in the mid-continental US: twelve years post-reflooding

In the mid-1980's, thousands of wetlands in the mid-continental Unites States were restored by interrupting drainage lines; revegetation of these systems, often cropped for decades and positioned in a predominantly agricultural landscape, relied solely on natural recolonization. A study of 64 of these wetlands determined that by 1991, three years after initial reflooding, aquatic species had efficiently recolonized whereas sedge meadow and wet prairie species had not. In 2000, 41 of these restorations that had not been significantly altered or returned to cultivation were revisited and their floras characterized by cover within distinct zones. While species richness increased on every site, the rate of accumulation varied widely. Furthermore, species that had colonized since 1991, including a variety of native wet prairie and sedge meadow species, were detected only at very low abundance. In contrast, Phalaris arundinacea L., an invasive perennial, was now present on every site, often at covers approaching 75–100% in the zones in which it occurred. Other invasive perennials, including Cirsium arvense (L.) Scop. and Typha angustifolia L./glauca Godr., had expanded significantly on many sites. The overall dominance of invasive perennials has resulted in basins that are becoming more similar over time. However, present variations in species richness and composition can be attributed to flooding frequency, and, potentially, basin size and isolation from nearby natural wetlands, as shown by TWINSPAN and graphical analysis. Basins that have not been flooded at midsummer for at least seven of 12 years are among the most depauperate in the study. Yet even frequently flooded basins lack diversity if they are small (< 1.5 ha) or isolated from seed sources. Across the study, numerous species common to natural systems were notably absent or infrequently occurring, including Calamagrostis canadensis (Michx.) Beauv., Carex lacustris Willd. and Lysimachia thrysiflora L. Continuous areas of sedge-dominated meadow, a signature of prairie potholes, did not exist at any study site, nor did they appear to be forming. Given the dominance of invasive perennials and absence of many native wetland species, it appears that without significant seeding, planting and aftercare wetland restorations in fragmented landscapes have a low probability of resembling those that existed historically. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of storage,mucilage presence,photoperiod, thermoperiod and salinity on germination of Farsetia aegyptia Turra (Brassicaceae) seeds: implications for restoration and seed banks in Arabian Desert

Seed viability and germination are key factors in the success of restoration efforts, especially when stored seeds are used. However, the effect of seed storage on germination of most of the native Arabian species is not well documented. We investigated the effect of storage time and role of the seed mucilage in regulating germination, dormancy, salinity tolerance and consequential survival strategy of F. aegyptia in an unpredictable arid desert setting. Effect of light and temperature during germination was studied under two photoperiods and two thermoperiods using intact and de-mucilaged seeds. Presence of mucilage and thermoperiod did not affect the germination. However, seed collection year and photoperiod had a highly significant effect on the germination. Increasing salinity levels decreased the germination of F. aegyptia but ungerminated seeds were able to germinate when salinity stress was alleviated. Seed storage at room temperature enhances the germination percentage, indicating that F. aegyptia seeds have physiological dormancy and it can be alleviated by after-ripening at dry storage. In addition, F. aegyptia seeds show ability to germinate at lower salinity concentration and remain viable even at higher saline conditions, indicating their adaptability to cope with such harsh environmental conditions.     

Response of Plant Height,Species Richness and Aboveground Biomass to Flooding Gradient along Vegetation Zones in Floodplain Wetlands,Northeast China

Flooding regime changes resulting from natural and human activity have been projected to affect wetland plant community structures and functions. It is therefore important to conduct investigations across a range of flooding gradients to assess the impact of flooding depth on wetland vegetation. We conducted this study to identify the pattern of plant height, species richness and aboveground biomass variation along the flooding gradient in floodplain wetlands located in Northeast China. We found that the response of dominant species height to the flooding gradient depends on specific species, i.e., a quadratic response for Carex lasiocarpa, a negative correlation for Calamagrostis angustifolia, and no response for Carex appendiculata. Species richness showed an intermediate effect along the vegetation zone from marsh to wet meadow while aboveground biomass increased. When the communities were analysed separately, only the water table depth had significant impact on species richness for two Carex communities and no variable for C. angustifolia community, while height of dominant species influenced aboveground biomass. When the three above-mentioned communities were grouped together, variations in species richness were mainly determined by community type, water table depth and community mean height, while variations in aboveground biomass were driven by community type and the height of dominant species. These findings indicate that if habitat drying of these herbaceous wetlands in this region continues, then two Carex marshes would be replaced gradually by C. angustifolia wet meadow in the near future. This will lead to a reduction in biodiversity and an increase in productivity and carbon budget. Meanwhile, functional traits must be considered, and should be a focus of attention in future studies on the species diversity and ecosystem function in this region.     

Increasing substrate heterogeneity as a bet‐hedging strategy for restoring wetland vegetation

Mimicking the natural heterogeneity of wetland substrates, e.g. by roughening surface soil or constructing hummocks, has been shown to facilitate wetland plant establishment. We asked if incorporating substrate heterogeneity could also help plants withstand variation in moisture levels. In a wetland with Carex stricta (tussock sedge) as the main restoration target, we manipulated substrates to create different soil moisture environments for planted C. stricta plugs. Our artificial mounds mimicked tussocks formed by C. stricta in natural meadows (circa 10–40 cm in height); we also varied mound compositions and created shallow depressions. Monitoring demonstrated variation in soil moisture among our treatments and natural differences in soil moisture between experimental blocks. Additionally, rainfall varied from severe drought in year 1 to extreme rainfall in year 2. Plug survival, flowering, cover, biomass, leaf length, and growth rate all varied with treatment, block, and/or year. Interactions among those factors were common. Planting plugs in shallow depressions exacerbated stress in a wet block during a wet year, causing low survival. Planting plugs in moisture‐retaining peat pots allowed them to survive and sustain growth even in a dry block during a dry year. We conclude that heterogeneous substrates can be used to hedge against environmental variability by widening the range of microsites available within a restoration site and thereby moderating stressful conditions in some areas.     

Clonal and genetic diversity of Carex moorcroftii on the Qinghai-Tibet plateau

Carex moorcroftii Falc. ex Boott is a rhizomatous clonal sedge dominating vast alpine steppe and meadow vegetations in the hinterland of the Qinghai-Tibet plateau. To reveal the genetic and clonal structure of this species, nine populations were investigated using ten inter-simple sequence repeat (ISSR) markers. As compared to other rhizomatous Carex species, C. moorcroftii had lower genetic diversity (H_s = 0.10) at population level and higher genetic differentiation (G_st = 0.66) and lower gene flow (N_m = 0.26) between populations. Clonal diversity in C. moorcroftii in terms of Simpson index (D = 0.65) was comparable to that in other clonal species while lower than that in Carex species from the arctic and subarctic areas. The ratio of clonal diversity to genetic variation in C. moorcroftii was closely correlated with latitude, enabling a speculation about the northern migration of this species on this plateau.     

Protocols for Use of Potamogeton perfoliatus and Ruppia maritima Seeds in Large‐Scale Restoration

Restoration of submerged aquatic vegetation from seed has been hampered by a lack of information on the appropriate conditions for collecting, processing, and storing seeds prior to dispersal. Seeds must be processed and stored under conditions that maintain seed viability, meet dormancy requirements, and prevent premature germination. This study examined the effects of collection date, processing technique, aeration, storage and induction temperature and salinity, and storage period on seed germination of two mesohaline aquatic species, Potamogeton perfoliatus and Ruppia maritima. Collection date and processing technique were significant factors affecting seed yield from donor populations. Seeds of both species remained viable and germinated best when stored at 4°C, and then exposed to freshwater induction conditions. However, their responses to other factors differed. Aeration during storage was necessary in order to maintain viability of P. perfoliatus seeds, whereas it was unnecessary for R. maritima seeds. Storage in freshwater at 4°C prevented germination of P. perfoliatus seeds, while high salinity during cold storage was necessary to minimize premature germination of R. maritima. Mean germination time of P. perfoliatus was dependent on storage salinity; in contrast, mean germination time of R. maritima seeds was dependent on induction salinity. These differences indicate that the methods required to produce large quantities of underwater plant seed amenable to large‐scale restoration efforts must be tailored to the specific requirements of individual species and must consider the range of processes from initial harvest through seed testing prior to field establishment.