Freshwater Plant Communities Research Articles

A comparative analysis of eDNA metabarcoding and field surveys: Exploring freshwater plant communities in rivers

While environmental DNA (eDNA) metabarcoding holds promise as a holistic approach to assess vegetation changes and community composition across diverse spatial and temporal scales, systematic investigations of its efficacy compared to conventional field surveys remain scarce in the literature. The present study explores the differences in plant diversity recovered from field surveys and captured with a multi-marker eDNA metabarcoding approach (two nrDNA ITS1 and ITS2, and two cpDNA rbcL and trnL) from river water samples. The eDNA metabarcoding approach retrieved 46 aquatic plants (hydrophytes and helophytes) and 245 terrestrial plants, compared to 24 and 127 species identified from field surveys. On average, eDNA samples collected immediately downstream of the survey sites recovered 43 % and 39 % of the aquatic and terrestrial species observed, respectively. Discrepancies were explained by differences in taxonomic resolution, the stochasticity of the retrieval of rare and elusive species, and the presence of reference sequences. We found a significant positive correlation between spatial and community distances at scales ranging from 2 to 9 km and identified turnover as the driving force of these differences. Metabarcoding demonstrated sensitivity to community changes and both approaches converge on a similar community structure. Interestingly, eDNA samples collected immediately upstream of the survey sites exhibited significant species overlap with the downstream samples (c. 100 m apart). Overall, our results demonstrate that within-site species mismatches between the methods are nonnegligible, and they question the use of eDNA for generating complete species lists at scales comparable to our field surveys (< 100-m transects). However, with adequate sampling and a multi-marker metabarcoding approach, eDNA has the potential to approximate catchment gamma diversity with less sampling effort than conventional surveys.

Morphological and Anatomical Differentiation of Potamogeton gramineus in Relation to the Presence of Invasive Species Elodea nuttallii: A Case Study from Vlasina Lake, Serbia.

Elodea nuttallii represents non-native and highly invasive species in Europe that significantly influence freshwater plant communities by decreasing the diversity of native species. This study aimed to determine whether the morphological and anatomical features of Potamogeton gramineus, a native species in Vlasina Lake, differ between sites where it coexists with E. nuttallii and those where E. nuttallii is not present. Environmental variables such as water depth, temperature, pH, conductivity, saturation, and O2 concentration were included in the analysis. Analyses were conducted on 32 morphological and anatomical features of P. gramineus collected from six sites within Vlasina Lake, comprising three sites where E. nuttallii was present and three sites where it was absent. The datasets containing morphometric and environmental variables underwent analysis using standard univariate techniques (Descriptive, ANOVA), Tukey's Honest Significant Difference (HSD) test, Student's t-test, and the Mann-Whitney U test, as well as multivariate statistical methods such as Canonical Discriminant Analysis (CDA). The results show the presence of morphological differentiation among P. gramineus individuals across the analyzed sites. These findings suggest that morphological and anatomical features, such as epidermis, mesophyll, palisade, and aerenchyma tissue thickness in floating leaves, number, length, width, and the surface area of stomata, as well as the width of submersed leaves and stem aerenchyma tissue thickness, effectively differentiate individuals that coexist with E. nuttallii and individuals that growth without its presence. Moreover, they indicate that P. gramineus exhibits a notable ability to modify its morphological traits in response to invasion.

Biogeography of freshwater plants in four major lakes in Kenya

Freshwater plants play a pivotal role in the health and functioning of freshwater ecosystems, and they also offer a variety of benefits to humans. Despite their importance, there is a significant gap in our understanding of the macroecology of freshwater plants in tropical regions, where biodiversity is typically high, and ecosystems face substantial pressures from human activities. To bridge this knowledge gap, we conducted extensive field surveys and applied statistical methods to understand the patterns of distribution, density, and diversity of freshwater plants and their relationship with abiotic components in four freshwater lakes in Kenya. We aimed to answer four key questions: (1) How many freshwater plant species occur in these lakes? (2) What proportion of these freshwater plant species are native and alien to Kenya? (3) What proportion of these species have a history of being invasive somewhere in the world, including in Kenya? (4) Which water and sediment characteristics are significantly associated with the presence, density, and diversity of freshwater plants in the lakes? We inventoried a total of 163 freshwater plant species, of which 114 were native to Kenya and 49 were alien. Online database searches revealed that 114 species had a history of invasion globally, with 22 species already invasive in Kenya. Our statistical analysis showed that the presence and density of plant species were positively correlated with pH and turbidity, but negatively correlated with soluble reactive phosphorus in both water and sediments, ammonia-nitrogen in sediments, and nitrate-nitrogen in water. Shannon-Wiener diversity of the plant species was positively correlated with ammonia-nitrogen in water and nitrate-nitrogen in sediments but negatively correlated with longitude and electrical conductivity of the water. Pielou’s evenness within the communities showed positive correlations with soluble reactive phosphorus in both water and sediments, ammonia-nitrogen in sediments, and nitrate-nitrogen in water but was negatively correlated with water turbidity and pH. The number of species with a history of invasion was positively correlated with longitude and electrical conductivity but negatively correlated with ammonia-nitrogen and nitrate-nitrogen in water. This extensive inventory not only enriches our understanding of the biodiversity within these freshwater plant communities but also highlights the potential risks posed by species with a history of being invasive. The findings regarding environmental variables linked to the presence, density, diversity, and invasive potential of these species underscore the complexity of freshwater ecosystems and the array of factors influencing the composition of freshwater plant communities.

Toward freshwater plant diversity surveys with eDNA barcoding and metabarcoding

AbstractProviding reliable, cost‐effective data on species distribution is critical to ensuring biodiversity conservation. However, many species may go unrecorded by conventional surveys, especially in aquatic environments. Environmental DNA (eDNA) barcoding and metabarcoding are alternative approaches that could complete biodiversity estimates based on species observations. While eDNA surveys are being standardized for some animal groups (e.g., fish and amphibians), research on eDNA approaches for freshwater plant communities is just starting to bear fruit. Here, we synthesized the 22 studies that used eDNA barcoding and metabarcoding to survey plant biodiversity in freshwater systems. We present evidence that contemporary aquatic plants (macrophytes) and terrestrial plants can be detected in water and surficial sediment eDNA from lakes and rivers. The phenology (e.g., senescence) of the target taxa strongly influences species detection. The main application of eDNA barcoding targets the monitoring of invasive macrophytes, and barcoding assays are available for 14 taxa. The metabarcoding approach shows a range of applications: the detection of rare macrophytes, catchment‐scale floristic inventories, and sediment fingerprinting. Barcodes on the plastid genome (cpDNA) are preferred for both approaches: matK and trnH‐psbA for barcoding, trnL, and rbcL for metabarcoding. The intergenic transcribed spacer 1 (ITS1) from the nuclear ribosomal DNA (nrDNA) was used for designing eDNA barcoding assays on five invasive macrophytes. In contrast, only three metabarcoding studies used the ITS1 or IST2 with newly designed primers. However, metabarcoding applications should routinely use a multi‐locus approach, combining cpDNA and nrDNA barcodes. Barcode combinations and existing primers need further testing on eDNA samples. We recommend using local barcode reference databases (BRDs), ideally self‐made, to circumvent taxonomic gaps and heterogeneous sequences in public BRDs. Finally, new technologies and developments like droplet digital PCR and hybridisation capture offer new perspectives for eDNA‐based biodiversity monitoring approaches.

Compositional breakpoints of freshwater plant communities across continents

Unravelling patterns and mechanisms of biogeographical transitions is crucial if we are to understand compositional gradients at large spatial extents, but no studies have thus far examined breakpoints in community composition of freshwater plants across continents. Using a dataset of almost 500 observations of lake plant community composition from six continents, we examined, for the first time, if such breakpoints in geographical space exist for freshwater plants and how well a suite of ecological factors (including climatic and local environmental variables) can explain transitions in community composition from the subtropics to the poles. Our combination of multivariate regression tree (MRT) analysis and k-means partitioning suggests that the most abrupt breakpoint exists between temperate to boreal regions on the one hand and freshwater plant communities harbouring mainly subtropical or Mediterranean assemblages on the other. The spatially structured variation in current climatic conditions is the most likely candidate for controlling these latitudinal patterns, although one cannot rule out joint effects of eco-evolutionary constraints in the harsher high-latitude environments and post-glacial migration lags after Pleistocene Ice Ages. Overall, our study supports the foundations of global regionalisation for freshwater plants and anticipates further biogeographical research on freshwater plant communities once datasets have been harmonised for conducting large-scale spatial analyses.

Invasion by an exotic grass species homogenizes native freshwater plant communities

Abstract A growing body of evidence has shown that biological invasions cause shifts in species composition of communities in space and time. Although biological invasions are considered a major driver of biotic homogenisation worldwide, most previous studies were conducted at small spatial scales and over short time periods, which may have underestimated the impacts of exotic species on native communities. Using a unique dataset of aquatic plants sampled in 235 sites over 12 years (2007–2010 and 2015–2019) in a large reservoir (Itaipu Reservoir; 1350 km2), we analysed how the invasion of a non‐native grass Urochloa arrecta affects the species richness, ecological uniqueness (i.e. local contribution to beta diversity—LCBD) and temporal β–diversity of native plant communities. From 3934 surveyed plant communities, U. arrecta was recorded in 2888 samples and it was absent from 1046 samples. Overall, species richness and ecological uniqueness of native plant communities were markedly lower in sites invaded than non‐invaded by U. arrecta. From 2007 to 2019, the ecological uniqueness of native plants was 60% lower in the invaded than non‐invaded sites. Whereas in invaded sites species loss was the dominant process driving native communities over time, in non‐invaded sites gain of new native species was the primary process underlying community trajectories. Moreover, comparing native plant communities before and after the invasion of U. arrecta, species richness, ecological uniqueness and species gains of native plant communities decreased, whereas species losses increased after the invasion of U. arrecta. Finally, the positive relationship between native biodiversity and precipitation was stronger in non‐invaded than invaded sites. Synthesis. Our findings provide comprehensive evidence that an invasive plant is decreasing the spatial and temporal β–diversity of native plant communities through declining species richness, rather than simply correlating with them. This suggests that U. arrecta is driving native plants to become less diverse and homogeneous after the invasion, both spatially and temporally. Our findings illustrate that at broad scales, aquatic plant communities may become increasingly homogeneous with the increasing number of biological invasion events taking place worldwide.

Temporal beta diversity of lake plants is determined by concomitant changes in environmental factors across decades

Abstract To comprehensively understand the impact of anthropogenic activities on biodiversity, we must understand how biodiversity has changed over time and what are the underlying processes. A growing body of evidence has shown that beta diversity reveals more about temporal changes in biodiversity compared with alpha diversity. Temporal beta diversity indicates, for example, degrees of change in species composition at single locations through time. We examined whether freshwater plant communities showed different patterns in temporal beta diversity in relation to concomitant changes in environmental conditions across decades. To do this, we used presence–absence data of lake plants for five decades (1940s–2010s) from southern Finland and calculated temporal beta diversity indices (TBI) for each lake between pairs of decades to the whole community, hydrophytes and helophytes. To get insights into possible processes behind the observed trends, we decomposed TBIs into beta diversity contributed by either temporal losses or temporal gains of species. We related TBIs and their loss and gain components to lake landscape position and changes in environmental variables. Based on comparisons of TBIs between the survey decade pairs, the temporal change in aquatic plant communities was modest through decades. Hydrophyte assemblages have changed more than helophyte assemblages. The main changes in temporal beta diversity occurred from the 1940s to the 1970s, when the gain of new species was the dominant process in the lakes throughout the landscape. Following that period, there was only modest changes, but from the 2000s to the 2010s, the dominant process was the loss of species. Temporal changes in environmental conditions played a key role in explaining the TBI. Our results showed that relying on only two survey points in time can result in limited knowledge of the ecological phenomenon under study and, for example, an exceptional year in terms of weather conditions can hinder detecting overall long‐term trends in compositional changes. Therefore, future studies should try to combine data from several decades to overcome the typical limitations of temporal information.

Species identity and diversity effects on invasion resistance of tropical freshwater plant communities

Biotic resistance mediated by native plant diversity has long been hypothesized to reduce the success of invading plant species in terrestrial systems in temperate regions. However, still little is known about the mechanisms driving invasion patterns in other biomes or latitudes. We help to fill this gap by investigating how native plant community presence and diversity, and the presence of native phylogenetically closely related species to an invader, would affect invader Hydrilla verticillata establishment success in tropical freshwater submerged plant communities. The presence of a native community suppressed the growth of H. verticillata, but did not prevent its colonisation. Invader growth was negatively affected by native plant productivity, but independent of native species richness and phylogenetic relatedness to the invader. Native plant production was not related to native species richness in our study. We show that resistance in these tropical aquatic submerged plant communities is mainly driven by the presence and biomass of a native community independent of native species diversity. Our study illustrates that resistance provided by these tropical freshwater submerged plant communities to invasive species contrasts to resistance described for other ecosystems. This emphasizes the need to include understudied systems when predicting patterns of species invasiveness and ecosystem invasibility across biomes.

Catchment properties and the photosynthetic trait composition of freshwater plant communities.

Unlike in land plants, photosynthesis in many aquatic plants relies on bicarbonate in addition to carbon dioxide (CO2) to compensate for the low diffusivity and potential depletion of CO2 in water. Concentrations of bicarbonate and CO2 vary greatly with catchment geology. In this study, we investigate whether there is a link between these concentrations and the frequency of freshwater plants possessing the bicarbonate use trait. We show, globally, that the frequency of plant species with this trait increases with bicarbonate concentration. Regionally, however, the frequency of bicarbonate use is reduced at sites where the CO2 concentration is substantially above the air equilibrium, consistent with this trait being an adaptation to carbon limitation. Future anthropogenic changes of bicarbonate and CO2 concentrations may alter the species compositions of freshwater plant communities.

An approach to quantitative plant–soil relationships of saltcedar woodlands throughout central and southeastern Iberian Peninsula (Spain)

Saltcedar woodlands are plant formations dominated by several species of the genus Tamarix (Tamaricaceae), representing the potential vegetation in saline, subsaline and wet environments under dry and semi-arid conditions. The broad ecological range of Tamarix communities leads to a heterogeneous floristic composition, with significant differences among habitats in terms of soil and vegetation. Some classification systems for Tamarix communities are only based on vegetation features and do not take any quantitative soil characteristics into account. Twelve Tamarix populations were selected under different ecological environments throughout central and southeastern Iberian Peninsula (Spain). Soil samples and vegetation inventories were collected over the course of 1 year to establish the plant–soil relationships based on constrained ordination analyses. The results showed that three different edaphic gradients were relevant to define the floristic composition of the Spanish saltcedar woodlands: a sodium–moisture gradient, a sulphate–magnesium gradient and a texture gradient. On the basis of these findings, we suggest a new classification system for Tamarix woodlands for the Mediterranean area based on plant–soil relationships. Three vegetation types have been proposed: hyperhalophilous, mesohalophilous and freshwater plant communities. Hyperhalophilous plant communities were characterised by soils with high E.C., high Na+ concentration, low soil moisture and high percentage of clay, being usually dominated by T. boveana and halophytes. Mesohalophilous plant communities had soils with high E.C., high Mg2+ and SO42− concentrations and high percentage of sand, being dominated by T. gallica with mesohalophilous and nitrophilous species. Finally, freshwater plant communities typically showed low E.C., low Na+ concentration and high soil moisture, being characterised by T. gallica with riparian and nitrophilous plants. Since the studied saltcedar woodland communities are notably dependent on soil salinity and moisture, the control of the human activities and hydrological alteration should be considered as a priority to contribute to the global preservation of the Tamarix woodlands.

Submerged freshwater plant communities do not show species complementarity effect in wetland mesocosms.

It is a generally accepted theory that ecological functions are enhanced with increased diversity in plant communities due to species complementarity effects. We tested this theory in a mesocosm study using freshwater submerged plant beds to determine if increasing species number caused overyielding and species complementarity. We applied a maximum of four species in the plant beds corresponding to the typical species number in natural freshwater plant beds. We found no clear effects of species number (1-4) on biomass production and thus no conclusive overyielding and complementarity effect. This may be explained by low species differentiation among the four species in plant traits relevant for resource acquisition in freshwater, or that other species interactions, e.g. allelopathy, were inhibiting overyielding. The existing knowledge on species complementarity in aquatic plant communities is sparse and inconclusive and calls for more research.

Multiproxy analysis of inception and development of the Lac‐à‐la‐Tortue peatland complex, St Lawrence Lowlands, eastern Canada

The initial conditions for the development of a large peatland complex in the St Lawrence Lowlands were reconstructed to increase the understanding of early development and expansion modes in this region. Peatland basin morphometry was identified by creating a model based on over 1500 existing peat depth measurements, and six cores were extracted along transects from a central (deepest) location towards the margins. C accumulation rates and ecohydrological conditions were reconstructed from plant macrofossils, testate amoeba assemblages and 14C chronologies. Luminescence dating was performed to better delineate the timing of dune stabilization in the area and potentially related climate changes. Shallow freshwater plant communities acted as nuclei for the development of a rich minerotrophic fen around 10 300 cal. a BP in the deepest part of a shallow depression at the surface of the St‐Maurice river delta. Peat inception was followed by the paludification of peripheral parabolic dune systems. Luminescence dating suggested dune stabilization between 11 500 and 10 900 years ago. The initial rich fen persisted until 9500 cal. a BP, and was replaced by a poor fen dominated by sedges as a result of a decrease in mineral nutrient influx from upland runoff. The shift to ombrotrophic conditions in the oldest section of Lac‐à‐la‐Tortue peatland started around 5150 cal. a BP. This major ecohydrological change coincides with those observed in several other peatlands in southern Québec. Variations in carbon and peat accumulation rates in both ombrotrophic and minerotrophic sectors appear to have been primarily controlled by hydroseral succession, peat‐forming vegetation, hydrological conditions, topography and fire activity. This study is the first to provide a quantification of the total carbon pool of a peatland complex in southern Québec at 6.39 Mt C, corresponding to a mean C mass per area of 96.9 kg C m−2 (σ = 50.60 kg C m−2).

Environmental filtering and competitive exclusion drive biodiversity‐invasibility relationships in shallow lake plant communities

Abstract Understanding the processes that influence the diversity of ecological communities and their susceptibility to invasion by exotic species remains a challenge in ecology. In many systems, a positive relationship between the richness of native species and exotic species has been observed at larger spatial (e.g., regional) scales, while a negative pattern has been observed at local (e.g., plot) scales. These patterns are widely attributed to (1) biotic interactions, particularly biotic resistance, limiting invasions in high‐diversity locations, producing negative local‐scale relationships, and (2) native and exotic richness covarying at larger spatial scales as a function of environmental conditions and heterogeneity, producing positive large‐scale relationships. However, alternative processes can produce similar patterns and need to be critically evaluated to make sound inferences about underlying mechanisms. We aggregated a large dataset of aquatic vegetation surveys from 1,102 Minnesota shallow lakes collected over 13 years to quantify spatial and temporal patterns of community composition. Using those data and additional information on environmental conditions we evaluated evidence for four distinct mechanisms that could drive patterns of native and exotic species richness: biotic resistance, competitive exclusion, environmental filtering and environmental heterogeneity. We found the classic pattern of a negative native‐exotic richness relationship at local scales and a positive relationship at lake scales. However, we found no evidence for local‐scale biotic resistance; instead, competitive exclusion by invasive species appeared to reduce native species richness after locations became invaded. Evaluating the influence of environmental filtering and heterogeneity, we found that native and exotic species occupied somewhat different niches. Invaders were less sensitive to environmental gradients and more tolerant of a wider range of conditions. This segregation of habitat preferences alone could produce a negative local native‐exotic richness relationship and a positive regional pattern without the involvement of biotic interactions. Synthesis. Our findings conflict with established expectations, which come from research predominantly conducted in terrestrial ecosystems. This illustrates the importance of explicitly evaluating underlying mechanisms in diversity‐invasibility research and for comparisons across different types of ecosystems. Identification of different drivers of diversity also has direct implications for decisions about management of freshwater plant communities.

Long-term changes in freshwater aquatic plant communities following extreme drought

Prolonged periods of floodplain drying are becoming increasingly common due to severe drought events and the effects of river regulation. Using long-term monitoring, we assessed changes in freshwater plant community structure and composition before (2000–2002) and after (2010–2013) an extended drying period that resulted in two formerly persistent and three seasonally inundated wetlands in south-eastern Australia remaining continuously dry for durations ranging from 4.7 to 9.4 years. Plant community composition and structure changed significantly between pre- and post-dry stages in all wetlands. These changes were characterised by significant reductions in the percentage cover of aquatic species and the loss of formerly dominant aquatic species—in particular, the herbaceous perennial species, Eleocharis acuta R.Br. and E. sphacelata R.Br., and the aquatic grass, Pseudoraphis spinescens (R.Br.) Vickery. Small areas of E. acuta began to re-establish in the second and third years of wetland refilling, whereas E. sphacelata and P. spinescens did not re-establish and the percentage cover of aquatic species overall remained significantly below 2000–2002 levels throughout the 2010–2013 period. These results lend empirical support to our understanding of floodplain dynamics and resilience, and in particular, the loss of dominant perennial aquatic species and establishment of opportunistic annual species following extended wetland drying.

Bistability of mangrove forests and competition with freshwater plants

Halophytic communities such as mangrove forests and buttonwood hammocks tend to border freshwater plant communities as sharp ecotones. Most studies attribute this purely to underlying physical templates, such as groundwater salinity gradients caused by tidal flux and topography. However, a few recent studies hypothesize that self-reinforcing feedback between vegetation and vadose zone salinity are also involved and create a bistable situation in which either halophytic dominated habitat or freshwater plant communities may dominate as alternative stable states. Here, we revisit the bistability hypothesis and demonstrate the mechanisms that result in bistability. We demonstrate with remote sensing imagery the sharp boundaries between freshwater hardwood hammock communities in southern Florida and halophytic communities such as buttonwood hammocks and mangroves. We further document from the literature how transpiration of mangroves and freshwater plants respond differently to vadose zone salinity, thus altering the salinity through feedback. Using mathematical models, we show how the self-reinforcing feedback, together with physical template, controls the ecotones between halophytic and freshwater communities. Regions of bistability along environmental gradients of salinity have the potential for large-scale vegetation shifts following pulse disturbances such as hurricane tidal surges in Florida, or tsunamis in other regions. The size of the region of bistability can be large for low-lying coastal habitat due to the saline water table, which extends inland due to salinity intrusion. We suggest coupling ecological and hydrologic processes as a framework for future studies.

The Grotta Grande of Scario (Salerno, Italy): Archaeology and environment during the last interglacial (MIS 5) of the Mediterranean region

Archeological and paleo-environmental researches carried on the Grotta Grande site illustrate the importance of a multidisciplinary approach among archeologists, palynologists and paleontologists. The archaeology, fauna, pollen and micro-charcoal recovered in two short sedimentary successions (trenches A, F) located close to the entrance of the cave are discussed. The cave opens directly on the Tyrrhenian Sea, 2 km from Scario (Salerno, Campania, Southern Italy). The morphology of the cave and sedimentary processes were controlled by eustatic fluctuations during the late Middle Pleistocene and the early Late Pleistocene. The sea repeatedly occupied the cave. The cave was frequented by humans of Middle Palaeolithic culture. Archaeological and faunal record of the two trenches can be positioned within the climatic fluctuation posterior to the warm interglacial MIS 5e peak. Pollen have been retrieved in the older series (b–c) of trench A, referable to MIS 7–6. The most important archaeological finds are the occurrence of structures that indicate the living space (trench F) together with hearths and lithic industry. The latter is characterised by the presence of the Levallois system and by the prevalence of sidescrapers among tools. The systematic use of limestone is observed only in trench F. Large and small mammal remains recovered within the two trenches show that in the neighbourhood of the site, a diversified assemblage occurred, including eleven large mammals, among which the hippo, the straight tusked elephant and the narrow-nosed rhino are noteworthy, and fourteen small mammal taxa, mainly rodents, with a significant amount of glirids. The fauna is indicative of a temperate, forested Mediterranean environment. Variation of faunal composition suggests that environments underwent some minor fluctuations towards cooler and/or drier landscapes. Pollen indicates vegetation organized in different belts, mainly Mediterranean evergreen forest/maquis and mixed forest with conifers and deciduous broadleaved trees. Fresh water plant communities are mainly represented in trench A. Some records of “Tertiary taxa” suggest stands of survival of these plants in the surroundings of the cave. Over the time span, the landscape become more open, due to some episodes of steppe vegetation spread caused by variation in temperature and humidity. Deposit of this age are rather uncommon even in the Italian peninsula, and therefore the integration of sedimentological, archaeological, faunal and palynological data provides an important piece of information to the puzzling reconstruction of the Late Pleistocene Mediterranean environments before the onset of the glaciation.

Effects of oil on internal gas transport, radial oxygen loss, gas films and bud growth in Phragmites australis.

Oil pollution of wetlands is a world-wide problem but, to date, research has concentrated on its influences on salt marsh rather than freshwater plant communities. The effects of water-borne light oils (liquid paraffin and diesel) were investigated on the fresh/brackish wetland species Phragmites australis in terms of routes of oil infiltration, internal gas transport, radial O(2) loss (ROL), underwater gas films and bud growth. Pressure flow resistances of pith cavities of nodes and aerenchyma of leaf sheaths, with or without previous exposure to oil, were recorded from flow rates under applied pressure. Convective flows were measured from living excised culms with oiled and non-oiled nodes and leaf sheaths. The effect of oil around culm basal nodes on ROL from rhizome and root apices was measured polarographically. Surface gas films on submerged shoots with and without oil treatment were recorded photographically. Growth and emergence of buds through water with and without an oil film were measured. Internodes are virtually impermeable, but nodes of senesced and living culms are permeable to oils which can block pith cavity diaphragms, preventing flows at applied pressures of 1 kPa, natural convective transport to the rhizome, and greatly decreasing ROL to phyllospheres and rhizospheres. Oil infiltrating or covering living leaf sheaths prevents humidity-induced convection. Oil displaces surface gas films from laminae and leaf sheaths. Buds emerge only a few centimetres through oil and die. Oil infiltrates the gas space system via nodal and leaf sheath stomata, reducing O(2) diffusion and convective flows into the rhizome system and decreasing oxygenation of phyllospheres and rhizospheres; underwater gas exchange via gas films will be impeded. Plants can be weakened by oil-induced failure of emerging buds. Plants will be most at risk during the growing season.

Stream mosses as chemically‐defended refugia for freshwater macroinvertebrates

Marine and terrestrial studies show that small, sedentary herbivores that utilize plants as both food and habitat can gain enemy‐free space by living on hosts that are chemically defended from larger, generalist consumers. Although large herbivores are increasingly recognized as important consumers of macrophytes in freshwater communities, the potential indirect effects of herbivory on plant‐associated macroinvertebrates have rarely been studied. Here, we show that the large, generalist consumers in a riverine system, Canada geese, Branta canadensis, and crayfish, Procambarus spiculifer, both selectively consumed riverweed, Podostemum ceratophyllum, over an aquatic moss, Fontinalis novae‐angliae, even though moss comprised 89% of the total plant biomass on riverine rocky shoals. Moss supported twice as many plant‐associated macroinvertebrates as riverweed, suggesting that it might provide a spatial refuge from consumption by these larger consumers. Bioassay‐guided fractionation of moss extracts led to the isolation of a C18 acetylenic acid, octadeca‐9,12‐dien‐6‐ynoic acid, that deterred crayfish feeding. In contrast to results with Canada geese and crayfish, both the amphipod Crangonyx gracilis and the isopod Asellus aquaticus consumed significant amounts of moss but rejected riverweed in laboratory feeding assays. Moreover, neither amphipod nor isopod feeding was deterred by the crude organic extract of Fontinalis, suggesting that these mesograzers tolerate or circumvent the chemical defenses that deterred larger consumers. Thus, herbivory by large, generalist herbivores may drive freshwater plant community structure towards chemically defended plants and favor the ecological specialization of smaller, less mobile herbivores on unpalatable hosts that represent enemy‐free space.

Dominant environmental factors in wetland plant communities invaded by Phragmites australis in East Harbor, Ohio, USA

Elevation, standing crop, disturbance and soil fertility often emerge from studies of freshwater plant communities as the dominant environmental factors determining both species richness and species composition. Few studies in North America have investigated the relationship between these factors and species abundance (standing crop) and species composition in the context of invasion by Phragmites australis. This study explores the influence of key abiotic and biotic variables on species abundance and composition across three Lake Erie wetlands differing in hydrology and Phragmites abundance in East Harbor, Ohio, USA. Standing crop for 92 species was related to standard sediment analyses, wave exposure, distance to shoreline, elevation, light interference, species density, and Phragmites standing crop in each of 95 1 × 1 m quadrats by using canonical correspondence analysis (CCA). Elevation (Axis I) and Phragmites standing␣crop-soil fertility (Axis II) explained 35.7 and 26.2%, respectively, of the variation in the species–environment relationships. Wave exposure was not a primary component of the first four canonical axes. Axis I was instrumental in describing species composition, separating wet meadow species from marsh species. Axis II was inversely related to species density for both wet meadow and marsh species. These findings generally support prevailing models describing the distribution of wetland plants along environmental gradients. Two discrepancies were noted, however: (1) species density was highest in the most sheltered sites and (2) wave exposure was directly associated with Phragmites standing crop-soil fertility gradient. The structural integrity of Phragmites stems, topographic heterogeneity and differential responses to anthropogenic disturbance may contribute to departure from prevailing multivariate models. This information has direct implications for local and regional wetland managers.

Chemical Defenses Promote Persistence of the Aquatic Plant Micranthemum umbrosum

Five of the most common macrophytes from an aquaculture facility with high densities of the herbivorous Asian grass carp (Ctenopharyngodon idella) were commonly unpalatable to three generalist consumers-grass carp and the native North American crayfishes Procambarus spiculifer and P. acutus. The rooted vascular plant Micranthemum umbrosum comprised 89% of the total aboveground plant biomass and was unpalatable to all three consumers as fresh tissues, as homogenized pellets, and as crude extracts. Bioassay-guided fractionation of the crude extract from M. umbrosum led to four previously known compounds that each deterred feeding by at least one consumer: 3,4,5-trimethoxyallylbenzene (1) and three lignoids: beta-apopicropodophyllin (2); (-)-(3S,4R,6S)-3-(3',4'-methylenedioxy-alpha-hydroxybenzyl)-4-(3'',4''-dimethoxybenzyl)butyrolactone (3); and (-)-hibalactone (4). None of the remaining four macrophytes produced a chemically deterrent extract. A 16-mo manipulative experiment showed that the aboveground biomass of M. umbrosum was unchanged when consumers were absent, but the biomass of Ludwigia repens, a plant that grass carp preferentially consumed over M. umbrosum, increased over 300-fold. Thus, selective feeding by grass carp effectively eliminates most palatable plants from this community and promotes the persistence of the chemically defended M. umbrosum, suggesting that plant defenses play critical yet understudied roles in the structure of freshwater plant communities.