Co-existence of three species of amphibious plants in relation to spatial and temporal variation: Investigation of plant responses

Plant responses in Ludwigia peploides (Kunth) Raven ssp. montevidensis (Spreng.) Raven and Marsilea mutica Mett., and the introduced Myriophyllum aquaticum (Vellozo) Verdc. were observed and related to co-existence of the species in a freshwater lagoon in the Sydney basin. Water levels vary in the lagoon, and plants of all three species grow readily from stem fragments on freely draining, waterlogged and submerged soils. In Marsilea, sporelings develop rapidly from ruptured sporocarps when wet. In Ludwigia, the seeds germinate in white and in red light, but not in darkness or far-red light. They germinate readily on soil and in water, floating or when held submerged. At 30°C, germination is rapid, at 40°C slower and less, and at 10°C nil, but seeds imbibed at 10°C and then exposed at 15°C have greater and more rapid germination than seeds maintained at 15°C. In relation to seasonal variation, plants of all three species grew at 10, 20, 30 and 40°C on freely draining, waterlogged and submerged soils, with the optimum temperature for growth apparently lower in Myriophyllum than in the other two species. In relation to water depth, plants of all three species grew when submerged to I m; the stems of Ludwigia and Myriophyllum elongating and reaching the water surface, and the petioles in Marsilea elongating bringing the leaf blades on to the surface, each within a week during summer. The floating leaf blades in Marsilea increase in size with depth of submergence of the subtending shoots. Mean relative growth rates (R?) under the most favourable experimental conditions were least in plants of Marsilea, the species occurring along more exposed parts of the shoreline in the lagoon. It was concluded that, while water levels in the lagoon were stable, differences in growth form between the species were probably more critical in their co-existence than quantitative differences in their growth: the floating leaves of Marsilea, being more resistant to wind and wave action than the floating shoots of Ludwigia and Myriophyllum, occupy the more exposed parts of the lagoon 'S edge. The most sheltered parts are occupied by the shoots of Myriophyllum which ride high on the water and overtop shoots of Ludwigia. Shoots of Ludwigia appear to be more resistant than those of Myriophyllum to removal and damage by wind and wave action, and achieve their greatest abundance on the water surface in situations of intermediate exposure to wind and wave action. Using Grubb's (1977) classification of niches, in Bushells Lagoon the three species appear to be primarily differentiated in habitat niche, less in regeneration niche and even less in phenological niche. In the lagoon, their habitat niches apparently overlap in water depth but are differentiated along a gradient of exposure.     

Interaction of vascular plants and vesicular-arbuscular mycorrhizal fungi across a soil moisture-nutrient gradient

Summary Abundance and distribution of vascular plants and vesicular-arbuscular mycorrhizal (VAM) fungi across a soil moisture-nutrient gradient were studied at a single site. Vegetation on the site varied from a dry mesic paririe dominated by little bluestem (Schizachyrium scoparium) to emergent aquatic vegetation dominated by cattail (Typha latifolia) and water smartweed (Polygonum hydropiperoides). Plant cover, VAM spore abundance, plant species richness, and number of VAM fungi represented as spores, had significant positive correlations with each other and with percent organic matter. The plant and VAM spore variables had significant negative correlations with soil pH and available Ca, Mg, P and gravimetric soil moisture. Using stepwise multiple regression, Ca was found to be the best predictor of spore abundance. Test for association between plant species and VAM fungal spores indicated that the spores of Glomus caledonium are associated with plants from dry, nutrient poor sites and spores of gigaspora gigantea are positively associated with plants occurring on the wet, relatively nutrient rich sites. Glomus fasciculatum was the most abundant and widely distributed VAM fungus and it had more positive associations with endophyte hosts than the other VAM fungi. We found no relationship between beta niche breadth of plant species and the presence or absence of mycorrhizal infection. However, our data suggest that some plant species may vary with respect to their infection status depending upon soil moisture conditions that may fluctuate seasonally or annually to favor or hinder VAM associations.     

扎龙湿地藻类优势种生态位及其种间联结性动态分析

为了解扎龙湿地藻类群落结构的季节特征和演替规律,对扎龙湿地藻类群落进行了分析,主要涉及藻类优势度、生态位、MFG功能群划分、生态位重叠值及联结系数分析。结果表明：(1)在检出的8门311种藻类中,夏季种类丰富度最高,全年优势种为狭形纤维藻和啮蚀隐藻。(2)优势种种类和密度季节性差异较大,春季以金藻-绿藻种类为主,夏季以绿藻-隐藻种类为主,秋季以隐藻-绿藻种类为主。(3)生态位宽度值与MFG功能群划分结合分析表明,优势种可分为3个大类别,不同种类对资源利用情况的差异较大,各季节广生态位种能较好的反映扎龙湿地水体环境的季节变化趋势。(4)生态位重叠值在不同季节上有较大差异,O_ik>0.6的占比分别为42.86%(春)、28.57%(夏)和25.00%(秋),表明春季优势种种间资源的竞争最为突出,种间竞争强于夏、秋两季。(5)种间联结性检验结果显示,扎龙湿地藻类优势种间总体联结上呈正关联关系,但是种间联结较为松散,种与种之间相对独立。研究表明,扎龙湿地藻类群落结构生态位测度均存在明显的季节波动,较好反映出水环境季节变化特征,可用来指示湿地水体环境的变化。     

The microdistribution of mayflies (Ephemeroptera) in myriophyllum beds in Pennington Creek,Johnston County,Oklahoma

I studied the population densities and the microdistributions of the four most abundant mayfly (Ephemeroptera) nymphs in Myriophyllum heterophyllum beds in Pennington Creek, Johnston County, Oklahoma, from May to December, 1977. The section of the stream studied has relatively constant flow rates and temperatures throughout the year due to continuous inflow from groundwater. This uniform physical environment allows the plant beds to grow throughout the year and provide a permanent habitat. The four mayflies are Tricorythodes fictus (Tricorythidae), Caenis delicata (Caenidae), Baetis favistriga and B. quilleri (Baetidae). All four species are of similar size and feed by scraping periphyton from the surface of the Myriophyllum. T. fictus and the two species of Baetis have overlapping microdistributions in the leafy, upper one-half of the plant bed. T. fictus and C. delicata have overlapping microdistributions in the lower portion of the plant bed and roots. Each species has similar population densities where their distributions overlap. There is no significant selection for position with respect to current flow in the Myriophyllum for any of the species. Food and space are always abundant, especially in the top one-half of the Myriophyllum. There are no major changes in these trends over the year.     

CONTRASTING STOMATAL RESPONSES TO VARIABLE SUNLIGHT IN TWO SUBALPINE HERBS

Gas exchange was measured in two subalpine herbs during alternating periods of sun and shade that simulated natural cloud patterns. Stomatal conductance (g) in the subalpine herb Helianthella quinquenervis was reduced by as much as 75% during 5-min shade periods that ranged in photosynthetic photon flux density (PPFD) from 100 to 1,100 μmol m^–2 sec^–1. In contrast, responses in g in another herb, Frasera speciosa, to fluctuations in sunlight were small, apparently due to slower stomatal responses. Based on an earlier hypothesis that water stress may strongly influence nonsteady state gas exchange responses to fluctuations in PPFD, net photosynthesis and g were measured in these herbs as seasonal water stress increased and compared with responses in irrigated plants. Stomatal conductance was relatively unresponsive to changes in PPFD in F. speciosa regardless of water stress. In contrast, substantial decreases in g occurred for H. quinquenervis during shade only when this species experienced water stress later in the season. Little response in g was measured early in the season or in irrigated plants. The seasonal shift in nonsteady state responses in H. quinquenervis from little response in g to shade when plants were nonstressed to more rapid reductions in g as water stress increased would maximize carbon gain early in the season when soil water was abundant while conserving water during periods of soil drought.     

Seed Bank Variation Along a Water Depth Gradient in a Subtropical Lakeshore Marsh,Longgan Lake,China

Although zonation patterns of the standing vegetation along a water depth gradient in wetlands have been well described, few studies have explored whether such patterns also occur in the seed bank. This study examined patterns of the seed bank along a water depth gradient in three vegetation types (submerged zone, floating-leaved zone, and emergent zone) of a subtropical lakeshore marsh, Longgan Lake, China. Submerged zone is the deepest water and never exposed its soil to air, and was dominant by submerged species; floating-leaved zone is waterlogged soil even during drawdown and was dominant by Nelumbo nucifera; emergent zone is rarely exceeded 1 m water depth during the wet season (summer and autumn), and the marsh soil was usually exposed to air during drawdown (winter and spring), and is dominant by Zizania latifolia, Polygonum hydropiper and Scirpus yagara. It was found that many species were ubiquitous in the seed bank. Frequency of distribution and densities of the dominant species, however, varied significantly from zone to zone. A total of 22 species was recorded in submerged zone, 20 in floating-leaved zone, and 29 in emergent zone. Germinated seedling density was 1,580, 8,994 and 20,424 seedlings m⁻² in submerged zone, floating-leaved zone, and emergent zone, respectively. Submerged and floating-leaved species were significantly abundant in the submerged zone, while the emergent species were found predominantly in the emergent zone. A fern species, Ceratopsis pterioides, was the most abundant seedling in seed banks from all three zones. A TWINSPAN dendrogram and CCA ordination diagram clearly showed separation of species among sites with the emergent zone being well separated from the submerged zone and floating-leaved zone, thus revealing marked zonal patterns in species distributions in the seed bank. This pattern of zonation reflected the pattern in the standing vegetation.     

Changes in submerged plants at the South end of Cayuga Lake following Tropical Storm Agnes

Tropical Storm Agnes produced increased suspended sediments and decreased water transparency at the southern end of Cayuga Lake for a sustained period during the early summer of 1972. The 1972 community of submerged vascular plants was severely affected. Standing crops were drastically reduced and the species composition of the plant community changed compared to 1970. In 1973, standing crop values had increased to 87% of those of 1970. However, shifts in spatial distribution had occurred and the community had further changed to one highly dominated by Myriophyllum sp. with the virtual exclussion of some previously abundant forms, especially Heteranthera dubia.     

Phenology, starch allocation, and environmental effects on Myriophyllum aquaticum

Seasonal biomass and starch allocation patterns were determined from natural populations of Myriophyllum aquaticum that were sampled monthly from January 2006 to December 2007 in Mississippi. Water temperature, water depth, light irradiance, light transmittance, pH, and conductivity were also recorded during biomass harvests. Overall, few significant relationships were observed between the environmental factors tested and seasonal biomass. Submersed shoot biomass was negatively related (p < 0.01) with water temperature. Stolons accounted for 40–95% of total biomass followed by emergent shoot, submersed shoot, and root biomass. Percent starch in plant tissues was positively related to water temperature. Starch allocation was greatest in stolons where up to 16.3% of total starch was stored. Submersed shoots stored 0.6–11.0% of total starch followed by emergent shoots (0.4–7%). The roots of M. aquaticum stored less than 3.8% of total starch throughout the study period. Reduced biomass and starch storage occurred from October to March in both 2006 and 2007. Management strategies for this species could utilize an integrated approach to exploit times of low energy reserves (fall and winter), or to remove emergent shoots to gain access to the stolons and other submersed tissues.     

Morphological adaptations of emergent plants to water flow: a case study with Typha angustifolia, Zizania latifolia and Phragmites australis

1. Water velocity plays an important role in shaping plant community structure in flowing waters although few authors have yet attempted to explain the adaptation of plants to flow. 2. We aimed to test two hypotheses, that: (i) some emergent macrophytes reconfigure their shoot distribution in fast currents and form clumps, and (ii) the shape and morphology of such clumps minimises drag caused by the current. The study focuses on three emergent macrophytes that co‐occur along a gradient of water velocity. 3. The species showed a clear zonation in response to water depth and current velocity. Phragmites australis occupied shallower and more slowly flowing water than Typha angustifolia and Zizania latifolia, which had similar preferences. 4. Both T. angustifolia and Z. latifolia shoots were more clumped at high velocity, whereas they were more randomly distributed at low flow or in stagnant water. Because of the low shoot density, water flowed more easily through T. angustifolia clumps, whereas Z. latifolia clumps had a high shoot density and large amounts of trapped litter, causing stagnant water in the centre of the clump. The clumps of Z. latifolia with a high density of shoots were longer and narrower than T. angustifolia clumps. Phragmites australis was less tolerant of flow than the other two species and large amounts of litter trapped in the clumps impaired flow. 5. The shoot distribution of both T. angustifolia and Z. latifolia is reconfigured at high flow and this minimises drag on the clumps.     

Effects of different macrophyte growth forms on sediment and P resuspension in a shallow lake

The effects of floating-leaved, submerged and emergent macrophytes on sediment resuspension and internal phosphorus loading were studied in the shallow Kirkkojärvi basin by placing sedimentation traps among different plant beds and adjacent open water and by sediment and water samples. All the three life forms considerably reduced sediment resuspension compared with non-vegetated areas. Both among submerged (Ceratophyllum demersum, Potamogeton obtusifolius, Ranunculus circinatus) and emergent (Typha angustifolia) plants, resuspension rate was on average 43% of that in the adjacent open water, while within floating-leaved plants (Nuphar lutea) the corresponding value was 87%. The effects of submerged and emergent vegetation increased in the course of the growing season together with increasing plant density. Among floating-leaved vegetation, such seasonal trend in resuspension effects was not observed. Compared with the non-vegetated area, floating-leaved, submerged and emergent plants reduced internal phosphorus loading on average by 21, 12 and 26 mg m⁻² d⁻¹, respectively. The effects of floating-leaved plants on resuspension-mediated internal phosphosrus loading were thus comparable to the effects of the other two life forms.     

Seasonal variation in terrestrial resource subsidies influences trophic niche width and overlap in two aquatic snake species: a stable isotope approach

Quantifying diet is essential for understanding the functional role of species with regard to energy processing, transfer, and storage within ecosystems. Recently, variance structure in the stable isotope composition of consumer tissues has been touted as a robust tool for quantifying trophic niche width, a task that has previously proven difficult due to bias in direct dietary analyses and difficulties in integrating diet composition over time. We used carbon and nitrogen stable isotope analyses to examine trophic niche width of two sympatric aquatic snakes, banded watersnakes Nerodia fasciata and black swamp snakes Seminatrix pygaea inhabiting an isolated wetland where seasonal migrations of amphibian prey cause dramatic shifts in resource availability. Specifically, we characterized snake and prey isotope compositions through time, space, and ontogeny and examined isotope values in relation to prey availability and snake diets assessed by gut content analysis. We determined that prey cluster into functional groups based on similarity of isotopic composition and seasonal availability. Isotope variance structure indicated that the trophic niche width of the banded watersnake was broader (more generalist) than that of the black swamp snake. Banded watersnakes also exhibited seasonal variation in isotope composition, suggesting seasonal diet shifts that track amphibian prey availability. Conversely, black swamp snakes exhibited little seasonal variation but displayed strong ontogenetic shifts in carbon and nitrogen isotope composition that closely paralleled ontogenetic shifts in their primary prey, paedomorphic mole salamanders Ambystoma talpoideum. Although niche dimensions are often treated as static, our results demonstrate that seasonal shifts in niche dimensions can lead to changes in niche overlap between sympatric species. Such short‐term fluctuations in niche overlap can influence competitive interactions and consequently the composition and dynamics of communities and ecosystems.     

Aquatic grazers reduce the establishment and growth of riparian plants along an environmental gradient

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The abundance of phytophilous invertebrates on different species of submerged macrophytes*

SUMMARY. 1. We tested Krecker's model (1939) which states that the abundance of invertebrates per unit macrophyte biomass varies with plant species and is higher on plants with finely dissected leaves than on plants with broad leaves. The abundance of invertebrates was measured in thirteen lacustrine macrophyte beds in southern Québec, Canada. The model was tested for the total abundance of invertebrates and for the abundances of Chironomidae, Cladocera, Cyclopoida, Gastropoda, Hydracarina, Ostracoda and Trichoptera. 2. More epiphytic invertebrates were found on the dissected Myriophyllum spp. than on the broad-leaved Potamogeton amplifolius Tuckerm, P. robbinsii Oakes and Vallisneria americana Michx. (P<0.01). More invertebrates were also found on P. amplifolius than on P. robbinsii or V. americana (P<0.01). The total abundance of invertebrates was not systematically related to the degree of plant dissection. 3. The abundances of Chironomidae, Cladocera, Cyclopoida, Gastropoda, Hydracarina, Ostracoda and Trichoptera varied on different plant species (P<0.01). Contrary to Krecker's hypothesis, however, macrophytes with finely dissected leaves (Ceratophyllum demersum and Myriophyllum spp.) did not in general support more invertebrates per unit plant biomass than plants with large leaves (Potamogeton amplifolius, P. robbinsii and Vallisneria americana).     

Net primary productivity and spatial distribution of vegetation in an alpine wetland,Qinghai-Tibetan Plateau

To initially describe vegetation structure and spatial variation in plant biomass in a typical alpine wetland of the Qinghai-Tibetan Plateau, net primary productivity and vegetation in relationship to environmental factors were investigated. In 2002, the wetland remained flooded to an average water depth of 25 cm during the growing season, from July to mid-September. We mapped the floodline and vegetation distribution using GPS (global positioning system). Coverage of vegetation in the wetland was 100%, and the vegetation was zonally distributed along a water depth gradient, with three emergent plant zones (Hippuris vulgaris-dominated zone, Scirpus distigmaticus-dominated zone, and Carex allivescers-dominated zone) and one submerged plant zone (Potamogeton pectinatus-dominated zone). Both aboveground and belowground biomass varied temporally within and among the vegetation zones. Further, net primary productivity (NPP) as estimated by peak biomass also differed among the vegetation zones; aboveground NPP was highest in the Carex-dominated zone with shallowest water and lowest in the Potamogeton zone with deepest water. The area occupied by each zone was 73.5% for P. pectinatus, 2.6% for H. vulgaris, 20.5% for S. distigmaticus, and 3.4% for C. allivescers. Morphological features in relationship to gas-transport efficiency of the aerial part differed among the emergent plants. Of the three emergent plants, H. vulgaris, which dominated in the deeper water, showed greater morphological adaptability to deep water than the other two emergent plants.     

Loopholes in the regulation of invasive species: genetic identifications identify mislabeling of prohibited aquarium plants

Numerous invasive aquatic species introductions can be traced to the aquarium trade. Many potentially harmful aquarium species may be difficult to identify based on morphology alone. As such, some prohibited or invasive species may be available for purchase if they are mislabeled as species without restrictions. Here we compare molecular identifications to internet vendors’ identifications for accessions of a popular genus of aquarium plants that are difficult to distinguish morphologically (Myriophyllum; watermilfoils). Specifically, we identified the extensive mislabeling of M. heterophyllum—an invasive species in the northeastern and western US. Furthermore, genotypes of M. heterophyllum found in our aquarium survey have also been found in invasive populations, suggesting their potential introduction through escape from aquaria, water gardens, or nurseries. Two additional taxa were sold under incorrect names. Finally, our survey revealed that Myriophyllum taxa present in the aquarium trade generally have poorly known distributions and ecologies, and therefore their invasive potential is unknown. Our study confirms that molecular identification methods can provide a valuable tool to survey commercial pathways for potentially harmful species that are otherwise difficult to identify.     

DISTRIBUTION OF DIURNAL ACID METABOLISM IN THE GENUS ISOETES

Previous studies have revealed a crassulacean acid metabolism-like pathway in the submerged aquatic Isoetes howellii (Isoetaceae). A survey of 11 other species in this largely aquatic genus showed all had diurnal fluctuations in titratable acidity and malic acid concentration. The magnitude of the diurnal fluctuation in titratable acidity varied from 119 (μeq g^–1 fresh weight) for I. bolanderi to 44 for I. engelmanni. Although these species were grown under similar conditions in a greenhouse, it is questionable whether or not any significance can be attached to observed species specific differences in magnitude of diurnal acid fluctuation. This is suggested by a single species comparison of plants in the field with greenhouse grown plants which indicated nearly as much variation in magnitude of acid fluctuations as observed across all species in the greenhouse. Much of this variation may be due to seasonal differences. Substrate appears to have very little effect on magnitude of acid fluctuations. Since this survey covered most of the range of the latitudinal, elevational, habitat, and morphological variation in the genus, it is likely that diurnal acid metabolism will prove to be common in the genus Isoetes.     

Herbivory in omnivorous fishes: effect of plant secondary metabolites and prey stoichiometry

1. Many animals that consume freshwater macrophytes are omnivorous (i.e., they include both plant and animal matter in their diet). For invertebrate omnivorous consumers, selection of macrophyte species depends partly on the presence of secondary metabolites in plants, plant carbon/nutrient balances and/or physical structure of plants. However, little is known about the mechanisms influencing consumption of macrophytes in aquatic vertebrates. 2. For two fish species, the omnivorous rudd (Scardinius erythrophthalmus) and herbivorous grass carp (Ctenopharyngodon idella), feeding preferences were determined in three choice experiments. We tested (i) whether the presence of secondary metabolites and macrophyte stoichiometry affects macrophyte species selection by fish, (ii) the importance of macrophyte stoichiometry by manipulating the macrophytes experimentally and (iii) the rate of herbivory when the most palatable macrophyte is offered simultaneously with a common animal prey. 3. In a choice experiment with five species of submerged macrophytes (Callitriche sp., Chara globularis, Elodea nuttallii, Myriophyllum spicatum and Potamogeton pectinatus), Myriophyllum was clearly consumed least by both fishes, which strongly correlated with the highest phenolic concentration of this macrophyte. Additionally, a significant negative relationship was found between consumption and C : N ratio of the five macrophytes. The two most consumed macrophytes also had the lowest dry matter concentration (DMC). 4. In a second choice experiment, the C : N ratio of the least (Myriophyllum) and most (Potamogeton) palatable plants was manipulated by growing the macrophytes under fertilised and unfertilised conditions and subsequently feeding them to rudd. The avoidance of consumption of the chemically defended Myriophyllum by rudd was partly alleviated by the lowered C : N ratio. 5. The third choice experiment showed that both fishes preferred animal prey (the amphipod Gammarus pulex) over the most palatable macrophyte (Potamogeton) when offered simultaneously. The C : N ratio of the amphipods was about half that of the lowest C : N ratio measured in the macrophytes. Consumption by the fishes could not clearly be related to C : P or N : P ratios of prey items in any of the experiments. 6. We conclude that omnivorous fish avoid macrophytes that are chemically defended. However, when these defences are only minor, stoichiometry (C : N ratio) in combination with DMC may be a determining factor for consumption by vertebrate facultative herbivores.     

A SURVEY FOR 1,3,6,7-TETRAHYDROXY-C-GLYCOSYLXANTHONES EMPHASIZING THE “PRIMITIVE” LEPTOSPORANGIATE FERNS AND THEIR ALLIES

A survey for 1,3,6,7-tetrahydroxy-C-glycosylxanthones of representative species within the primitive vascular plants, emphasizing the leptosporangiate ferns, has indicated a limited distribution of these compounds within three leptosporangiate families: Hymenophyllaceae, Aspleniaceae and Marsileaceae. In the Hymenophyllaceae the distribution of these compounds appears to be a useful criterion for segregating species of Mecodium from other species of Hymenophyllum (sensu lato) and suggests that the tubulate vs. the valvate indusial condition may not be an ideal character for separating all species of Hymenophyllum (s.l.) from those of Trichomanes (s.l.). These compounds appear useful for delimiting several species of Elaphoglossum section Pachyglossa and support a relationship among the Aspleniaceae, Athyriaceae, and Elaphoglossaceae. Their presence in Marsilea also raises questions as to the origin of this group of plants.     

Seasonal dynamics of resource partitioning among foliage-gleaning passerines in northern Finland

Summary The five most abundant species were included in a year-round study with respect to six foraging niche dimensions. Approximately full multidimensional utilization functions were used for niche metrics. During summer foraging overlaps were invariably high, but in other seasons periodically lower. Foraging site breadth was lower in winter, when fewer sites are profitable for foraging than in summer. Feeding posture versatility, by contrast, was highest in winter. Seasonal foraging shifts were very prominent, as great in fact as between-species differences. Often seasonal trends were parallel in different species. Niche axes of macrohabitat-type (e.g. tree) were more open for foraging variation and axes of microhabitat-type (tree part) more rigid. Among the resident species seasonal variation in foraging was greatest in Regulus regulus and Parus cristatus, whereas the foraging behaviour was more stable for P. montanus (an abundant species with broad niche), perhaps owing to greater intraspecific competition. In these northern forests foliage-gleaners must be versatile generalists to cope with the unpredictable resources, and thus they overlap broadly in their general resource niches which are determined by their genetically fixed cost and benefit relations to each resource type. Anyhow, presumably during periodical food shortage, the actual resource uses adjusted by resource availability and competition may overlap narrowly.