Disentangling the effects of drought,salinity, and sulfate on baldcypress growth in a coastal plain restored wetland

Because of their dominance in swamps of the southeastern United States, baldcypress (Taxodium distichum) trees are commonly used in wetland restoration. Though baldcypress are known to tolerate moderate flooding and salinity, their growth has been shown to decrease when they experience drought and high salinity. This study examined the effects of drought and elevated salinity on the growth of baldcypress seedlings and saplings. In a restored wetland in North Carolina, we examined the growth of 8‐year‐old baldcypress by measuring height and diameter at breast height (DBH) along salinity, nutrient, and flooding gradients. In a greenhouse, we placed 1‐year‐old baldcypress seedlings in either drought or saturated conditions and applied different water treatments: fresh water, sulfate, and artificial salt water (5 ppt [parts per thousand]). Over 26 weeks, we measured diameter at root collar, height, and biomass. In the field, chloride concentrations in soil solution had a negative effect on DBH and height (51 and 36% decrease, respectively); high water levels had a negative effect on height (47% decrease) and DBH (46% decrease). In the greenhouse, both drought and salinity decreased diameter growth (43 and 61% decline, respectively) and height (64 and 43% decline, respectively). Sulfate did not have a significant effect on diameter growth, but caused a 14% decrease in height. Our results suggest that both drought and salinity (even as low as 1 ppt) can lead to a 20–60% decline in baldcypress growth. Restoration practitioners should consider the negative consequences of both drought and increased salinity on baldcypress growth when planning for wetland restoration.     

Flooding and salinity effects on growth and survival of four common forested wetland species

The survival, growth, and biomass of baldcypress (Taxodium distichum (L.) Rich.), water tupelo (Nyssa aquatica L.), Chinese tallow (Sapium sebiferum (L.) Roxb.), and green ash (Fraxinus pennsylvanica Marsh.) seedlings were examined in an experiment varying water levels (watered, flooded) and salinity levels (0, 2, and 10 ppt, plus a simulated storm surge with 32 ppt saltwater). All seedlings, except for those flooded with 10 ppt saltwater, survived to the end of the experiment. In 10 ppt saltwater, flooded baldcypress, water tupelo, and green ash survived two weeks whereas Chinese tallow survived for 6 weeks. However, a second set of slightly older baldcypress, water tupelo, and Chinese tallow seedlings survived eight weeks of flooding with 10 ppt saltwater. When carried through the winter to the beginning of the second growing season, flooded baldcypress and Chinese tallow seedlings from the 0 and 2 ppt treatments leafed out, but only Chinese tallow recovered from the saltwater surge treatment. The diameter and growth (height) of each species was not affected when watered with 2 ppt saltwater, except for the effects of the height growth of baldcypress. Growth was reduced for all species when watered with 10 ppt saltwater. The diameter growth of green ash was reduced by freshwater flooding. The diameter growth of baldcypress and water tupelo was greater when flooded with fresh water. Flooding with 2 ppt saltwater caused a significant reduction in diameter growth in water tupelo, green ash, and Chinese tallow, but not in baldcypress. Root and stem biomass values were not significantly different for any species between the 0 and 2 ppt salinity watering treatments. However, seedlings watered with 10 ppt saltwater had significantly lower root and stem biomass values, except for baldcypress roots and green ash stems. Baldcypress was least affected by flooding with 0 and 2 ppt saltwater, although there were slight reductions in root biomass with increasing salinity. Because of the susceptibility of the seedlings of these four species to increases in flooding and salinity, their regeneration may be limited in the future, thereby causing shifts in species composition.     

Effects of Flooding on Susceptibility of Taxodium distichum L. Seedlings to Drought

Responses of baldcypress (Taxodium distichum) seedlings to soil moisture were studied to test the hypothesis that flooding may lead to seedling's higher susceptibility to drought. Treatments included a well-watered but drained control (C), continuously flooded (CF), control followed by drought (CD), and flooded followed by drought (FD). Gas exchange values revealed no significant effects on net photosynthetic rate (P _N) in response to flooding. In contrast, after the onset of drought, P _N was significantly reduced in CD and FD plants. Significant growth reductions under mild drought conditions indicated that baldcypress seedlings were drought sensitive. However, comparison of gas exchange rates and growth responses between CD and FD plants indicated that prior flooding had no detectable effect on subsequent sensitivity of baldcypress to drought. These findings explain baldcypress persistence in wetland habitats characterized by periodic flooding and mild drought.     

Effects of flood pre-conditioning on responses of three bottomland tree species to soil waterlogging

The potential for improving the performance of seedlings of woody species under flood stress was evaluated. Seedlings of baldcypress (Taxodium distichum), nuttall oak (Quercus nuttallii), and swamp chestnut oak (Quercus michauxii) were subjected to a two-phase study in which soil moisture regime was controlled. During Phase I, plants were randomly assigned to either: Control, well-watered and drained conditions; or intermittent flooding, alternating flooding and drained conditions. Following completion of Phase I, seedlings in each treatment were assigned randomly to one of two treatments: well-watered but drained or continuously flooded. Thus during Phase II, plants were divided into four groups: Control (A), intermittent flooding during Phase I and draining during Phase II (B), control during Phase I and continuously flooding during Phase II (C), and intermittently flooded during Phase I and continuously flooded during Phase II (D). Stomatal conductance (gw) in baldcypress increased in flooded treatments ranging from 112 to 128 percnt; of controls. Net photosynthesis (Pn) in baldcypress was not significantly affected by flooding while in oaks, Pn and gw decreased significantly in response to flooding. In oaks, flood pre-conditioning did not affect growth or physiological responses during phase II significantly. However, stomatal conductance of flood-hardened baldcypress was increased compared to seedlings that were not hardened. The study species developed significantly higher root porosity under flooded conditions compared to controls. Flood pre-conditioning did not appear to increase root porosity although it may have accelerated the process.     

RESPONSE OF WOODY SWAMP SEEDLINGS TO FLOODING AND INCREASED WATER TEMPERATURES. I. GROWTH,BIOMASS, AND SURVIVORSHIP

Growth, biomass, and survival of bald cypress (Taxodium distichum [L.] Richard), water tupelo (Nyssa aquatica L.), black willow (Salix nigra Marshall), and button bush (Cephalanthus occidentalis L.) were examined in a 3 times 3 factorial experiment varying water temperatures (AMBIENT, MID, and HIGH [~40 C]) and water levels (DRAINED, SATURATED, and FLOODED). Stem diameter and height, biomass, and survivorship for water tupelo and bald cypress were all reduced by the HIGH/FLOODED treatment. Black willow growth had the greatest variability among nonlethal flooding and temperature treatments, and achieved the greatest biomass of the four species. In the HIGH/FLOODED treatment, however, only 47% of the black willow seedlings survived and stem diameter, height, and biomass of survivors were greatly reduced. Button bush had intermediate variability of growth to the nonlethal treatments as compared to the other study species. Survival of button bush seedlings in the HIGH/FLOODED treatment was high (87%), but root biomass of the survivors was reduced. Interspecific differences in growth, biomass, survivorship, and morphological characteristics existed among these swamp species to experimental conditions. These responses may help explain vegetation patterns in a thermally impacted swamp.     

Partial flooding enhances aeration in adventitious roots of black willow (Salix nigra) cuttings

Black willow (Salix nigra) cuttings are used for streambank stabilization where they are subjected to a range of soil moisture conditions including flooding. Flooding has been shown to adversely impact cutting performance, and improved understanding of natural adaptations to flooding might suggest handling and planting techniques to enhance success. However, data assessing the root aeration in adventitious roots that are developed on cuttings of woody species are scant. In addition, it appears that no data are available regarding aeration of the root system under partially flooded conditions. This experiment was designed to examine the effects of continuous flooding (CF) and partial flooding (PF) on aerenchyma formation and radial oxygen loss (ROL) in black willow cuttings. Photosynthetic and growth responses to these conditions were also investigated. Under laboratory condition, replicated potted cuttings were subjected to three treatments: no flooding (control, C), CF, and PF. Water was maintained above the soil surface in CF and at 10 cm depth in PF. Results indicated that after the 28-d treatments, root porosity ranged between 28.6% and 33.0% for the CF and C plants but was greater for the PF plants (39.2% for the drained and 37.2% for the flooded portions). A similar response pattern was found for ROL. In addition, CF treatment led to decreases in final root biomass and root/shoot ratio. Neither CF nor PF had any detectable adverse effects on plant gas exchange or photosystem II functioning. Our results indicated that S. nigra cuttings exhibited avoidance mechanisms in response to flooding, especially the partially flooded condition which is the most common occurrence in riparian systems.     

Effects of soil moisture regimes on photosynthesis and growth in cattail (Typha latifolia)

Both waterlogging and water deficiency are major environmental factors affecting plant growth and functioning in many wetland and floodplain ecosystems across North America. Wetland plants possess various characteristics that enable them to survive and function in the intermittently flooded wetland environments, while their sensitivity to drought has received less attention. The present study quantified the photosynthetic and growth responses of cattail (Typha latifolia), an important species of freshwater wetlands, to a wide range of soil moisture regimes. In addition, changes in the efficiency of photosynthetic apparatus following initiation of the treatments were investigated. Under greenhouse conditions, seedlings were subjected to four soil moisture regimes: (1) drained (control), (2) continuous flooding, (3) periodic flooding, and (4) periodic drought. Results indicated that dark fluorescence yield was increased in response to periodic drought, while it showed decreases under continuous flooding. Net photosynthesis and stomatal conductance were enhanced by continuous flooding and periodic flooding. In contrast, these parameters exhibited reduction under periodic drought. In addition, leaf chlorophyll content was adversely affected by periodic drought. Recovery of net photosynthesis was noted, along with enhanced height growth, in both continuously and periodically flooded plants. Meanwhile, continuous flooding enhanced biomass production while periodic drought led to biomass reduction. Periodic drought also contributed to substantial reduction in root growth compared with shoot growth. Therefore, the combined photosynthetic performance and growth responses of cattail are likely to contribute to the ability of this species to thrive in flooded condition but be susceptive to periodic drought.     

Effects of Pre-Planting Soaking on Growth and Survival of Black Willow Cuttings

Black willow (Salix nigra) uses periodic flood events for dispersal of vegetative propagules, subjecting them to periods of soaking before their deposition along the streambank. It was hypothesized that this life history trait results in optimal conditions for willow growth and survival. To test this hypothesis, a greenhouse experiment was conducted using 1.2‐m‐long black willow cuttings (posts) with a basal diameter of 5 cm. Cuttings were subjected to three soaking treatments (0, 3, and 10 days) and then grown under four soil moisture regimes (control, drought, permanently flooded, and intermittently flooded). Growth, biomass, and survival were recorded. Results showed that soaking posts for 10 days was most beneficial in the control soil moisture regime, enhancing root, shoot, leaf, and total biomass production. Shoot abundance and growth were also enhanced by 10 days of soaking in the control and permanently flooded moisture regimes. Finally, survival increased significantly in control and drought moisture regimes in response to the 10‐day soaking treatment. Results clearly demonstrated that soaking had significant effects on willow post success when evaluated across all soil moisture regimes. Posts subjected to the 10‐day soaking treatment consistently had greatest growth and biomass and displayed a doubling of the survival rate. Therefore, soaking willow cuttings before planting is a simple, inexpensive technique that may be used to bolster streambank restoration success.     

Is there evidence of adaptation to tidal flooding in saplings of baldcypress subjected to different salinity regimes?

Plant populations may adapt to environmental conditions over time by developing genetically based morphological or physiological characteristics. For tidal freshwater forested wetlands, we hypothesized that the conditions under which trees developed led to ecotypic difference in response of progeny to hydroperiod. Specifically, we looked for evidence of ecotypic adaptation for tidal flooding at different salinity regimes using growth and ecophysiological characteristics of two tidal and two non-tidal source collections of baldcypress (Taxodium distichum (L.) L.C. Rich) from the southeastern United States. Saplings were subjected to treatments of hydrology (permanent versus tidal flooding) and salinity (0 versus ～2 g l^?1) for two and a half growing seasons in a greenhouse environment. Saplings from tidal sources maintained 21–41% lower overall growth and biomass accumulation than saplings from non-tidal sources, while saplings from non-tidal sources maintained 14–19% lower overall rates of net photosynthetic assimilation, leaf transpiration, and stomatal conductance than saplings from tidal sources. However, we found no evidence for growth or physiological enhancement of saplings from tidal sources to tide, or of saplings from non-tidal sources to no tide. All saplings growing under permanent flooding exhibited reduced growth and leaf gas exchange regardless of source, with little evidence for consistent salinity effects across hydroperiods. While we reject our original hypothesis, we suggest that adaptations of coastal baldcypress to broad (rather than narrow) environmental conditions may promote ecophysiological and growth enhancements under a range of global-change-induced stressors, perhaps reflecting a natural resilience to environmental change while precluding adaptations for specific flood regimes.     

Responses of three bottomland species with different flood tolerance capabilities to various flooding regimes

Seedlings of baldcypress (Taxodium distichum), nuttall oak (Quercus nuttalli), and cherrybark oak (Quercus falcata var.pagodaefolia) were subjected to four flooding treatments: control, continuously flooded, intermittently flooded, and partially flooded for 70 days in a greenhouse. The treatments imposed various durations and intensities of soil redox potential (Eh) conditions representing a range encountered by plants in their habitats. Morphological changes and gas exchange responses to the treatments differed among the study species. Rapid development of adventitious root and hypertrophied lenticels were observed in baldcypress and nuttall oak under all flooded treatments. Cherrybark oak had the highest percentage reduction in net photosynthesis ranging from 65–87%, whereas reductions in nuttall oak ranged between 35–68% and in baldcypress between 6–21% in response to various treatments. Recovery of gas exchange was noted in baldcypress but no significant recovery was found in oaks. The recovery in baldcypress contributed to the continued growth and biomass accumulation under various treatments. Little evidence of consistent changes in biomass allocation patterns in response to the treatments was found in baldcypress but total biomass decreased significantly under the continuously flooded treatment. In oaks, total biomass decreased significantly in all flooded treatments. The present findings demonstrated that physiological functions are adversely affected by low soil Eh conditions and the extent of such effects are dependent on the intensity and duration of soil reduction as well as the species' capability to respond to such conditions rapidly. Management plans concerned with regeneration of bottomland forested ecosystems should consider the species flood response capabilities at seedling stages as well as the timing, durations, and intensities of soil reduction at the specific site.     

Regeneration potential of <Emphasis Type="Italic">Taxodium distichum</Emphasis> swamps and climate change

Seed bank densities respond to factors across local to landscape scales, and therefore, knowledge of these responses may be necessary in forecasting the effects of climate change on the regeneration of species. This study relates the seed bank densities of species of Taxodium distichum swamps to local water regime and regional climate factors at five latitudes across the Mississippi River Alluvial Valley from southern Illinois to Louisiana. In an outdoor nursery setting, the seed banks of twenty-five swamps were exposed to non-flooded (freely drained) or flooded treatments, and the number and species of seeds germinating were recorded from each swamp during one growing season. Based on ANOVA analysis, the majority of dominant species had a higher rate of germination in non-flooded versus flooded treatments. Similarly, an NMS comparison, which considered the local water regime and regional climate of the swamps, found that the species of seeds germinating, almost completely shifted under non-flooded versus flooded treatments. For example, in wetter northern swamps, seeds of Taxodium distichum germinated in non-flooded conditions, but did not germinate from the same seed banks in flooded conditions. In wetter southern swamps, seeds of Eleocharis cellulosa germinated in flooded conditions, but did not germinate in non-flooded conditions. The strong relationship of seed germination and density relationships with local water regime and regional climate variables suggests that the forecasting of climate change effects on swamps and other wetlands needs to consider a variety of interrelated variables to make adequate projections of the regeneration responses of species to climate change. Because regeneration is an important aspect of species maintenance and restoration, climate drying could influence the species distribution of these swamps in the future.      

Flooding effects on rapid responses of the invasive plant Alternanthera philoxeroides to defoliation

In experiments under controlled growth conditions it was examined how flooding affected the responses of the invasive plant Alternanthera philoxeroides to defoliation. In drained and flooded conditions, plants were subjected to five defoliation levels: 0, 10, 50, 90% removal of leaf tissue and apex removal (90% leaf tissue plus apical bud removal). Plants were harvested weekly for five weeks. In drained conditions, plant biomasses including total biomass, shoot biomass and root biomass after 50% defoliation rapidly recovered to the control plant level. They were significantly lower for the 90% defoliation and apex removal treatments compared to control plants throughout the experiment. In flooded conditions, total biomass and shoot biomass after 50% defoliation, 90% defoliation, and apex removal treatments could return to control plant levels before the end of the experiment. In 90% defoliation and apex removal treatments root to shoot biomass ratios of both drained and flooded plants were initially much higher than in control plants, but the difference disappeared rapidly. The final biomasses decreased with increased defoliation intensity in drained conditions, but no significant difference was generally found in any of the defoliation treatments in flooded conditions. The rapid re-growth of A. philoxeroides plants after defoliation may partly be responsible for its invasion success. However, defoliation capable of removing 90% of the leaf tissue may be desirable in restricting the growth of this invasive species in drained conditions.     

Responses of red alder and black cottonwood seedlings to flooding

Red alder ( Alnus rubra Bong.) and black cottonwood ( Populus trichocarpa Torr. & Gray) seedlings were monitored to evaluate response during a 20-day period of artificial flooding and a 20-day recovery period following flooding. During the flooding period, both species showed changes in nutrient uptake and transport, initiated stemderived adventitious roots that became aerenchymatous, and exhibited hypertrophied lenticels. Flooded red alder seedlings also showed reduced height and leaf area growth and developed lower-stem hypertrophy. Flooded black cottonwood seedlings exhibited root dieback, aerenchyma in below ground root tips, and changes in root hydraulic conductance and xylem pressure potential. Contrary to expectations, however, stomatal closure following flooding was not observed in either species. Flooded red alder seedlings increased growth rapidly when drained, and by the end of the recovery period, formerly flooded and non-flooded red alder seedlings differed only minimally in this respect. In contrast, several characteristics of black cottonwood – including growth rate and nutrient content – still differed between formerly flooded and non-flooded seedlings at the end of the recovery period. Based on observed treatment differences at the end of the experiment, red alder seedlings were judged to be more tolerant of flooding than black cottonwood.     

Growth rates of baldcypress (Taxodium distichum) seedlings in a treated effluent assimilation marsh

Wetlands have proven effective at improving water quality of treated wastewater effluent, which in turn promotes increased primary productivity and vertical accretion. Baldcypress (Taxodium distichum) seedlings grown under different conditions (bare root and potted) were planted in four subunits of an effluent assimilation marsh and a control marsh in southeast Louisiana, USA, and basal diameter growth was monitored over one growing season. Mean basal diameter growth for seedlings in the assimilation subunits ranged from 16.1 (±1.4) mm to 9.5 (±0.9) mm, whereas growth for seedlings planted in the control marsh was 6.4 (±0.9) mm. Seedlings planted nearest the outfall experienced greater basal diameter growth (18.1 ± 2.6) compared to those planted 700 m away (8.0 ± 0.9), with growth generally decreasing with distance. Potted seedlings experienced greater growth (19.1 ± 1.0 and 20.6 ± 1.0 for five-month-olds and ten-month olds, respectively) than bare root seedlings (4.6 ± 0.6 and 4.0 ± 0.4 for one-year-olds and two-year olds, respectively). Planting assimilation marshes with baldcypress seedlings can be an effective restoration tool for coastal Louisiana, which will provide hurricane protection and improved surface water quality. Wastewater treatment wetlands may offer an effective tool for restoring coastal baldcypress (T. distichum)-water tupelo (Nyssa aquatic) swamps in Louisiana.     

Comparative studies on physiological and biochemical adaptation of Taxodium distichum and Taxodium ascendens seedlings to different soil water regimes

Responses of baldcypress (Taxodium distichum) and pondcypress (Taxodium ascendens) seedlings in leaf gas exchange and root metabolism to a wide range of hydrological regimes were studied. Four water treatments included control (C), mild drought (MD), wet soil (WS) and flooding (FL). Both species showed no significant change in net photosynthetic rate (Pn) in response to flooding in contrast to significant reduction in Pn under mild drought conditions. In baldcypress, Pn was also maintained at control level compared to significant increase of Pn in pondcypress when subjected to the wet soil treatment. In flooding, both species showed significant increase in contents of malate and shikimate in their lateral roots and further in total roots, as compared to control. However, different adaptation to flooding was demonstrated in their taproots in contents of malate and shikimate. Furthermore, baldcypress seedlings showed significant reductions in lateral and total root biomass, in contrast with no significant change in that of pondcypress seedlings in response to flooding. In wet soil, no significant effects were revealed in contents of malate and shikimate in roots (including different root portions) of baldcypress seedlings in contrast to significant increase of their contents in lateral roots of pondcypress seedlings. No significant effects on root biomass were detected in either baldcypress or pondcypress seedlings in response to wet soil. When subjected to mild drought, baldcypress seedlings displayed no significant change in contents of malate and shikimate in roots, whereas pondcypress seedlings exhibited significant reduction in content of shikimate in lateral roots and total roots. In addition, root biomass of baldcypress seedlings showed no significant change, while lateral and total root biomass of pondcypress seedlings demonstrated significant increase when mild drought was imposed. These findings indicate that baldcypress and pondcypress appear to be promising candidates for reforestation in the Three Gorges Reservoir region due to their characteristics of water-tolerance and mild drought endurance. However, pondcypress is more resilient than baldcypress to the continuum of water availability conditions expected in the Three Gorges Reservoir region.     

Physiological responses of black willow (<Emphasis Type="Italic">Salix nigra</Emphasis>) cuttings to a range of soil moisture regimes

We quantified the physiological responses of black willow to four soil moisture regimes: no flooding (control, C), continuous flooding (CF), periodic flooding (PF), and periodic drought (PD). Stomatal limitation was one of the factors that led to the reduced photosynthetic capacity in CF cuttings. Under PD, stomatal closure, decreased leaf chlorophyll content, and increased dark fluorescence yield contributed to photosynthetic decline. CF cuttings accumulated the lowest shoot biomass while the final height and root growth were most adversely affected by PD. PF cuttings tended to allocate more photoassimilates to root growth than to shoots.     

Involvement of plant growth substances in the alteration of leaf gas exchange of flooded tomato plants

Ethylene, abscisic acid, and cytokinins were tested for their ability to either induce or prevent the changes which occur in gas exchange characteristics of tomato (Lycopersicon esculentum Mill. cv. Rheinlands Ruhm) leaves during short-term soil flooding. Ethylene, which increases in the shoots of flooded plants, had no effect on stomatal conductance or photosynthetic capacity of drained plants. Abscisic acid, which also accumulates in the shoots of flooded plants, could reproduce the stomatal behavior of flooded plants when sprayed on the leaves of drained plants. However, photosynthetic capacity of drained plants was unaffected by abscisic acid sprays. Cytokinin export from the roots to the shoots declines in flooded plants. Spray applications of benzyladenine increased stomatal conductance in both flooded and drained plants. In addition, the decline in photosynthetic capacity during flooding was largely prevented by supplementary cytokinin applications. The possible involvement of these growth substances in modifying leaf gas exchange during flooding is discussed.     

Connecting carbon and nitrogen storage in rural wetland soil to groundwater abstraction for urban water supply

We investigated whether groundwater abstraction for urban water supply diminishes the storage of carbon (C), nitrogen (N), and organic matter in the soil of rural wetlands. Wetland soil organic matter (SOM) benefits air and water quality by sequestering large masses of C and N. Yet, the accumulation of wetland SOM depends on soil inundation, so we hypothesized that groundwater abstraction would diminish stocks of SOM, C, and N in wetland soils. Predictions of this hypothesis were tested in two types of subtropical, depressional‐basin wetland: forested swamps and herbaceous‐vegetation marshes. In west‐central Florida, >650 ML groundwater day^?1 are abstracted for use primarily in the Tampa Bay metropolis. At higher abstraction volumes, water tables were lower and wetlands had shorter hydroperiods (less time inundated). In turn, wetlands with shorter hydroperiods had 50–60% less SOM, C, and N per kg soil. In swamps, SOM loss caused soil bulk density to double, so areal soil C and N storage per m² through 30.5 cm depth was diminished by 25–30% in short‐hydroperiod swamps. In herbaceous‐vegetation marshes, short hydroperiods caused a sharper decline in N than in C. Soil organic matter, C, and N pools were not correlated with soil texture or with wetland draining‐reflooding frequency. Many years of shortened hydroperiod were probably required to diminish soil organic matter, C, and N pools by the magnitudes we observed. This diminution might have occurred decades ago, but could be maintained contemporarily by the failure each year of chronically drained soils to retain new organic matter inputs. In sum, our study attributes the contraction of hydroperiod and loss of soil organic matter, C, and N from rural wetlands to groundwater abstraction performed largely for urban water supply, revealing teleconnections between rural ecosystem change and urban resource demand.     

With a little help from my friends: physiological integration facilitates invasion of wetland grass Elymus athericus into flooded soils

Tidal wetlands worldwide are undergoing rapid invasions by tall-growing clonal grasses. Prominent examples are invasions by species of the genera Spartina, Phragmites and Elymus. The responsible physiological and ecological drivers of these invasions are poorly understood. Physiological integration (PI) is a key trait of clonal plants, which enables the exchange of resources among ramets. We investigated PI in Elymus athericus, which has been rapidly spreading from high-marsh into low-marsh environments of European salt marshes during the last decades. We applied a nitrogen stable-isotope approach to trace nutrient translocation between ramets in a factorial mesocosm experiment. The experiment was set up to mimic an invasion pattern commonly found in tidal wetlands, i.e. from high-elevated and rarely flooded into low-elevated and frequently flooded microenvironments. We tested for intraspecific variability in PI by including two genotypes of Elymus that naturally occur at different elevations within the tidal frame, a high-marsh (HM) and a low-marsh (LM) genotype. PI strongly increased offspring ramet aboveground and belowground biomass by 62 and 81%, respectively. Offspring ramets under drained conditions had 95% greater belowground biomass than those under flooded conditions. LM genotype offspring ramets produced 27% more aboveground biomass than HM genotypes. Offspring ramets were clearly more enriched in ¹⁵N under flooded versus drained conditions; however, this positive effect of flooding on δ¹⁵N was only significant in the LM genotype. Our findings demonstrate the importance of PI for the growth of Elymus offspring ramets and thereby for the species' capacity for fast vegetative spread. We show that offspring ramets under stressful flooded conditions are more dependent on nutrient supply from parent ramets than those under drained conditions. Our data furthermore suggest a higher degree of adaptation to flooding via PI in the LM versus HM genotype. In conclusion, we highlight the importance of assessing PI and intraspecific trait variability to understand invasion processes within ecosystems.     

Establishment and Early Growth of Willow at Different Levels of Weed Competition and Nitrogen Fertilization

To evaluate the effects of weed competition and nitrogen fertilization on the early growth performance of willow, cuttings of the clone Tora (Salix schwerinii x S. viminalis) were planted in buckets together with model weeds (spring barley or white mustard) sown 15, 26, and 30 days after willow planting. The buckets were fertilized with 30 or 90 kg N ha^?1. Willow with weeds sown after 15 days produced less biomass and smaller leaf area and had a lower maximum shoot height compared to willow planted without weeds and willow with weeds sown after 26 or 30 days. Fertilization with 90 kg N ha^?1 gave higher willow biomass production in willow with weeds sown after 26 or 30 days. Type of model weed had no effect on willow performance. Weed biomass and maximum shoot height were higher in weeds planted without willows compared to the willow-weed mixtures. A high nitrogen level gave more weed biomass when planted without willows and in the willow-weed mixture with weeds sown after 15 days. We conclude that for the given high density of willow, competition from weeds emerging soon after willow planting had strong effect on early production. Furthermore, if there is a risk of weed infestation, fertilization should be delayed.