Spatial pattern of native species Myriophyllum spicatum and invasive alien species Elodea nuttallii after introduction of the latter one into the Drava River (Slovenia)

Invasive alien species Elodea nuttallii was introduced into the Drava River (Slovenia) in 2007. The spatial distribution of native macrophyte species Myriophyllum spicatum and the invasive alien species E. nuttallii was studied in two impoundments, HPP Vuhred and HPP Mariborski otok in the years 2009, 2010 and 2011. The very heterogeneous environment of the Drava River had resulted in dynamic and non-uniform distributions of E. nuttalli and M. spicatum in both impoundments. The distribution of E. nuttalli was shown to be suppressed in river stretches exposed to great turbulence. It had, however, successfully invaded areas with gentle slope of littoral, low exposition to main water flow and muddy sediment and prevailed against the native species M. spicatum. The latter was more abundant than E. nuttallii in stretches exposed to higher flow velocity. Thus, the competitive success of M. spicatum and E. nuttallii depended on environmental conditions. Our results lead to the prediction that E. nuttallii will increase further in the river Drava, causing serious economic problems.     

Merging geospatial and field data to predict the distribution and abundance of an exotic macrophyte in a large Wisconsin reservoir

Geospatial and field data were merged to characterize the invasion of an exotic macrophyte into a large (∼6192 ha), reservoir, the Chippewa Flowage, Wisconsin, USA. Multi-scale analyses provided data for quantitative models of human, abiotic and biotic influences that would explain the presence and abundance of Myriophyllum spicatum L., a widespread, aquatic species. Human and abiotic factors were analysed in a geographic information system and additional abiotic and biotic factors were sampled at 2778 sample points. Using multivariate logistic and multiple linear regressions the relationships between 29 factors and the species’ presence and abundance were quantified. M. spicatum occurred at 181 of 992 vegetated points. M. spicatum abundance was significantly influenced by two geospatial variables (wave action and water-level drawdown) and two site variables (water depth and native species presence). Separate models for shallow and deep water explained 32% and 30% of M. spicatum abundance, respectively (compared to 22.8% using all 181 points). The species’ vertical distribution (range = 0.30-4.27 m) was related to depth of winter drawdown (∼1.3 m on average) and to water depth (an indicator of decreasing light), as well as to shading by floating-leaved macrophytes. Spatial extrapolation of model predictions for M. spicatum presence indicated that a total 940 hectares are at risk for M. spicatum invasion (∼15% of the Chippewa Flowage; about 2.5 times larger than the current distribution).     

Allelopathic inhibition of epiphytes by submerged macrophytes

The hypothesis that epiphytes are more vulnerable to allelochemicals released by submerged macrophytes than phytoplankton was tested by measuring growth and photosystem (PS) II activity of three common epiphytic algae and cyanobacteria in coexistence with Myriophyllum spicatum using dialysis tubes. Results were compared with earlier experiments on planktonic species. Contrary to the planktonic species, the tested epiphytes, the green algae Stigeoclonium tenue, the diatom Gomphonema parvulum and the cyanobacterium Oscillatoria limosa, were not significantly inhibited by M. spicatum. Growth and PS II activity of O. limosa were even significantly enhanced by M. spicatum, but this effect disappeared under phosphorus-deficiency due to the allelopathically induced inhibition of the alkaline phosphatase activity or phosphorus leakage by the macrophytes. My findings of a lower vulnerability of epiphytes against allelopathic substances of submerged macrophytes are supported by results of a literature survey.     

Chemical profile of the North American native Myriophyllum sibiricum compared to the invasive M. spicatum

Myriophyllum spicatum L. is a nonindigenous invasive plant in North America that can displace the closely related native Myriophyllum sibiricum Komarov. We analyzed the chemical composition (including: C, N, P, polyphenols, lignin, nonpolar extractables, and sugars) of M. spicatum and M. sibiricum and determined how the chemistry of the two species varied by plant part with growing environment (lake versus tank), irradiance (full sun versus 50% shading), and season (July through September). M. spicatum had higher concentrations of carbon, polyphenols and lignin (C: 47%; polyphenols: 5.5%; lignin: 18%) than M. sibiricum (C: 42%; polyphenols: 3.7%; lignin: 9%) while M. sibiricum had a higher concentration of ash under all conditions (12% versus 8% for M. spicatum). Apical meristems of both species had the highest concentration of carbon, polyphenols, and tellimagrandin II, followed by leaves and stems. Tellimagrandin II was present in apical meristems of both M. spicatum (24.6 mg g⁻¹ dm) and M. sibiricum (11.1 mg g⁻¹ dm). Variation in irradiance from 490 (shade) to 940 (sun) μmol of photons m⁻² s⁻¹ had no effect on C, N, and polyphenol concentrations, suggesting that light levels above 490 μmol of photons m⁻² s⁻¹ do not alter chemical composition. The higher concentration of polyphenols and lignin in M. spicatum relative to M. sibiricum may provide advantages that facilitate invasion and displacement of native plants.     

Programmed cell death in the cyanobacterium <Emphasis Type="Italic">Microcystis aeruginosa</Emphasis> induced by allelopathic effect of submerged macrophyte <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> in co-culture system

This investigation demonstrates that programmed cell death (PCD) in a cyanobacterium, Microcystis aeruginosa, resulting from allelopathic stress induced by a submerged macrophyte, Myriophyllum spicatum, in a co-culture system. The hallmarks of PCD, caspase-3-like protease activity, DNA fragmentation, and destruction of cell ultrastructure, as well as intracellular PCD signaling radicals, reactive oxygen species (ROS), and nitric oxide (NO), were measured in M. aeruginosa cells co-cultured with M. spicatum for 7 days. The results showed a dose–response relationship between M. spicatum biomass and M. aeruginosa mortality. A caspase-3-like protease was activated and elevated from day 3. Thylakoid disintegration, cytoplasmic vacuolation, and fuzzy nuclear zone were observed by transmission electron microscopy, and distinct DNA fragmentation was detected in M. aeruginosa cells at a M. spicatum biomass of 6.0 g fresh weight (FW) L^?1 during the 7 days. Allelochemicals of total phenolic compounds (TPCs) were determined in co-culture water, and the concentration increased with increasing of M. spicatum biomass and co-culture time. Compared with the level of ROS production in the control group, a significant overproduction of ROS was detected in M. aeruginosa cells in the treatment group, and this was positively correlated with TPC concentration. Furthermore, the level of intracellular NO increased with the percent mortality of M. aeruginosa. The results indicated that a PCD pathway was induced in the cyanobacterium M. aeruginosa when co-cultured with the submerged macrophyte M. spicatum.     

Food web interactions determine energy transfer efficiency and top consumer responses to inputs of dissolved organic carbon

The sand stored in the gizzard of some freshwater snails should assist in the mechanical digestion of tough food, yet effects of gizzard sand on consumption of living macrophyte tissue are seldom addressed. We quantified the effects of gizzard sand both on consumption of plant tissue and on snail growth and reproduction with a cross-gradient, 28-day laboratory experiment. We offered apical shoot sections of Myriophyllum spicatum (initial ash content ~ 56%) or soft lettuce (ash content ~ 19%, as control) to Lymnaea stagnalis previously reared on sand or without sand. Gizzard sand had no effect on snail growth but sand-reared snails fed with lettuce initially produced more eggs than their no-sand counterparts. Snails fed M. spicatum exhibited stunted growth and produced only one small egg clutch. Shell height at time of first oviposition of M. spicatum-fed snails was two-thirds of that of lettuce-fed controls. Snails initially appeared to feed on periphyton and older, brownish M. spicatum leaflet tips, but later turned to midstem tissue. Fresh apices were not consumed. Gizzard sand had a limited effect on the ability of L. stagnalis to consume living M. spicatum, with apices likely defended chemically (polyphenols) and midstem tissue defended structurally (high ash content).     

Fertile sediment and ammonium enrichment decrease the growth and biomechanical strength of submersed macrophyte Myriophyllum spicatum in an experiment

Decline of submersed macrophytes has occurred in eutrophic lakes worldwide. Little is known about effects of nutrient enrichment on biomechanical properties of submersed macrophytes. In a 30-day experiment, Myriophyllum spicatum was cultured in aquaria containing two types of sediment (mesotrophic clay vs. fertile loam) with contrasting water NH₄ ⁺ concentrations (0 vs. 3.0 mg L^?1 NH₄–N). The plant growth, shoot and root morphology, stem biomechanical properties, and stem total nonstructure carbohydrates content (TNC) were examined. The NH₄ ⁺-enriched water, particularly combined with the fertile sediment, caused adverse effects on M. spicatum as indicated by reductions in the growth, stem biomechanical properties (tensile force, bending force and structural stiffness), and TNC content. These results indicate that increased sediment fertility and water NH₄ ⁺-enrichment made the plant more fragile and vulnerable to hydraulic damage, particularly for the upper stem, implying that M. spicatum was prone to uprooting and fracture by hydraulic force, and the broken fragment from parent shoot of M. spicatum might have low-survival potential due to its low-TNC content. This may be a mechanical aspect for the decline of submersed macrophytes and makes it more difficult to restore submersed vegetation in the eutrophic lakes.     

The effect of temperature, water level and burial depth on seed germination of Myriophyllum spicatum and Potamogeton malaianus

The effects of temperature, water level and burial depth on seed germination of two submerged species, Myriophyllum spicatum and Potamogeton malaianus, were investigated under controlled laboratory conditions. There was no significant difference in final germination of M. spicatum among water level treatments, but P. malaianus germinations at 1 cm and 12 cm water levels were better than at 0 cm water level at temperatures of 20 °C and 30 °C. Little to no germination was observed for either species at the temperature of 10 °C. At 15 °C, however, germination increased significantly to 66.3-70.6% for M. spicatum and to 29.4-48.1% for P. malaianus under all three water level treatments. Increased temperature from 15 °C to 30 °C had no significant effect on the final germination of M. spicatum except at the 1 cm water level, but enhanced significantly the germination of P. malaianus. Analysis of the mean time to germination revealed that M. spicatum was a faster germinator relative to P. malaianus. The two species’ germination differed markedly in response to burial depth. Germination percentage of M. spicatum was 71.3% at 0 cm burial depth, but decreased to 5.0% and to 2.5% at depths of 1 cm and 2 cm, respectively; whereas germination percentages of P. malaianus were 40.0%, 23.8%, 12.5%, 7.5% and 1.3% at depths of 0 cm, 1 cm, 2 cm, 3 cm and 5 cm, respectively. We concluded that the two species respond differently to germination strategies. The findings provided further insight into how germination strategy contributes to the seed bank formation and species invasion.     

Resprouting Response of Aquatic Clonal Plants to Cutting May Explain Their Resistance to Spate Flooding

Resprouting ability may increase a plant??s resistance to recurrent disturbances in aquatic ecosystems. We investigated the effect of mechanical disturbances on survival and regrowth patterns in three clonal aquatic species of similar growth form but with different ecological ranges in terms of flooding (Myriophyllum verticillatum, Myriophyllum spicatum and Potamogeton coloratus). P. coloratus prefers to colonize stable habitats, whereas M. verticillatum occurs in intermediately flooded habitats and M. spicatum is tolerant to a high flooding frequency. Two cutting treatments (single cuts or repeated cuts) were applied under controlled conditions. We hypothesized that M. verticillatum and M. spicatum would be resistant to cutting displaying either a tolerant or an escape strategy whereas P. coloratus would be sensitive to cutting. Our hypothesis was validated, as the three species displayed contrasting responses to disturbance. M. verticillatum displayed efficient clonal propagation following breakage (escape strategy), but its growth rate decreased after recurrent disturbances. P. coloratus displayed a close response but was unable to compensate biomass loss even after one cut. M. spicatum maintained a similar growth rate by developing a densely branched form despite recurrent disturbances but with a low investment in clonal growth (tolerance strategy). Both biomass compensation and clonal propagation influence plant fitness, but their relative advantage differs depending on the flooding frequency experienced by plants in their natural habitats. Clonal propagation may promote recolonization after disturbances in infrequently flooded sites, but seems less efficient than a tolerance strategy for survival in frequently flooded sites.     

Expansion of Myriophyllum spicatum (Eurasian water milfoil) into Lake Nasser,Egypt: Invasive capacity and habitat stability

The invasion of Myriophyllum spicatum into Lake Nasser, and its impact on submerged macrophyte communities are quantitatively documented. Samples of macrophytes, water and hydrosoil were collected from 17 sites, in October and November 2002. The average dry weight standing crop of each species per grapnel haul was determined at each depth zone (sampling site). Twenty-one environmental variables were measured (12 water and 9 hydrosoil variables). Canonical correspondence analysis (CCA) was used to determine species–environment relationships. Comparing these relationships of the present study with those detected in 1988–1990 indicated significant changes in water and hydrosoil characteristics. These changes are also implicated in the submerged macrophyte communities. M. spicatum has replaced the originally dominant submerged macrophyte Najas marina subsp. armata. The study indicated that the invasion of M. spicatum depends not only on its attributes, but also on the physico-chemical characteristics of Lake Nasser.     

Experimental estimation of the possible use of submersed macrophytes for biotesting bottom sediments of the Yenisei River

The laboratory testing of bottom sediments (BSs) from the Yenisei River containing different concentrations of technogenic radionuclides, heavy metals, and biogenic elements (N and P) based on aquatic such plants as Elodea canadensis (Canadian waterweed) and Myriophyllum spicatum (Eurasian watermilfoil) has revealed a higher sensitivity of roots to the general quality of BSs than shoots: shoot length (9%) < root length (11%) < root number (15%) in M. spicatum; shoot length (22%) < root length (42%) < root number (44%) in E. canadensis. In contrast to M. spicatum, the growth parameters of roots and shoots in E. canadensis have differed in a significant statistical manner between most BS samples. A reverse correlation has been found between the increase in shoot length and the activity of technogenic radionuclides in BSs, which is mostly significant in E. canadensis (r ² = 0.90–0.95, p = 0.05). Since the growth of shoots and roots in E. canadensis has turned out to be more sensitive to changes in the quality of BSs than that in M. spicatum, E. canadensis can be considered more prospective for biotesting BSs.     

Experimental Tests of Priority Effects and Light Availability on Relative Performance of Myriophyllum spicatum and Elodea nuttallii Propagules in Artificial Stream Channels

Submersed macrophytes have important ecological functions in many streams, but fostering growth of beneficial native species while suppressing weedy invasives may be challenging. Two approaches commonly used in management of terrestrial plant communities may be useful in this context: (1) altering resource availability and (2) establishing desirable species before weeds can invade (priority effects). However, these approaches are rarely used in aquatic systems, despite widespread need for sustainable solutions to aquatic weed problems. In artificial stream channels in California, USA, I conducted experiments with asexual propagules of non-native invasive Myriophyllum spicatum (Eurasian watermilfoil) and native Elodea nuttallii (western waterweed) to address the questions: (1) How does light availability affect relative performance of the two species?; (2) Does planting the native earlier than the invasive decrease survival or growth rate of the invasive?; and (3) Do light level and priority effects interact? The relative performance between E. nuttallii and M. spicatum had an interesting and unexpected pattern: M. spicatum had higher growth rates than E. nuttallii in the zero and medium shade levels, but had similar performance in the low and high shade levels. This pattern is most likely the result of E. nutallii’s sensitivity to both very low and very high light, and M. spicatum’s sensitivity to very low light only. Native priority did not significantly affect growth rate or survival of M. spicatum, possibly because of unexpectedly poor growth of the E. nuttallii planted early. This study suggests that altering light levels could be effective in reducing growth of an invasive macrophyte, and for changing the competitive balance between a native and a non-native species in the establishment phase. Further investigations into the use of priority effects and resource alteration for submersed macrophyte management are warranted, given their mixed results in other (limited) studies.     

Enumeration and Characterization of Bacterial Colonists of a Submersed Aquatic Plant, Eurasian Watermilfoil (Myriophyllum spicatum L.)

A simple procedure for enumerating and grouping the bacterial colonists of Eurasian watermilfoil (Myriophyllum spicatum L.) is described. Colony characteristics of bacteria associated with M. spicatum were better defined and more stable on nutrient-poor, diluted nutrient broth agar than on high-nutrient media. Acinetobacter, Cytophaga, Flavobacterium, Moraxella, Pseudomonas and/or Alcaligenes, and Vibrio/Aeromonas spp., as well as two highly fastidious unidentified bacterial groups (gram-negative rods and gram-negative cocci), were associated with cultured watermilfoil during January, February, May, June, July, and August 1988. In Lake Wingra (Madison, Wis.), Micrococcus spp. and enterobacters were also associated with Eurasian watermilfoil during July, August, and October 1987.     

Reproductive Allocation in Three Macrophyte Species from Different Lakes with Variable Eutrophic Conditions

Reproductive allocation is a key process in the plant life cycle and aquatic plants exhibit great diversity in their reproductive systems. In the present study, we conduct a field investigation of three aquatic macrophytes: Stuckenia pectinata, Myriophyllum spicatum, and Potamogeton perfoliatus. Our results showed that widespread species, including S. pectinata and M. spicatum had greater plasticity in their allocation patterns in the form of increased sexual and asexual reproduction, and greater potential to set seeds and increase fitness in more eutrophic environments. P. perfoliatus also exhibited a capacity to adopt varied sexual reproductive strategies such as setting more offspring for the future, although only in clear conditions with low nutrient levels. Our results establish strategies and mechanisms of some species for tolerating and surviving in varied eutrophic lake conditions.     

Behavioural thermoregulation by the endangered crocodile lizard (Shinisaurus crocodilurus) in captivity

Understanding the factors that may affect behavioural thermoregulation of endangered reptiles is important for their conservation because thermoregulation determines body temperatures and in turn physiological functions of these ectotherms. Here we measured seasonal variation in operative environmental temperature (T_e), body temperature (T_b), and microhabitat use of endangered crocodile lizards (Shinisaurus crocodilurus) from a captive population, within open and shaded enclosures, to understand how they respond to thermally challenging environments. T_e was higher in open enclosures than in shaded enclosures. The T_b of lizards differed between the open and shaded enclosures in summer and autumn, but not in spring. In summer, crocodile lizards stayed in the water to avoid overheating, whereas in autumn, crocodile lizards perched on branches seeking optimal thermal environments. Crocodile lizards showed higher thermoregulatory effectiveness in open enclosures (with low thermal quality) than in shaded enclosures. Our study suggests that the crocodile lizard is capable of behavioural thermoregulation via microhabitat selection, although overall, it is not an effective thermoregulator. Therefore, maintaining diverse thermal environments in natural habitats for behavioural thermoregulation is an essential measure to conserve this endangered species both in the field and captivity.     

Uptake of stable N isotopes by Myriophyllum spicatum is not selective

Preference for ¹⁴N over ¹⁵N was assessed for Myriophyllum spicatum L. by exposing shoots to the same initial concentration of N, but different isotopic compositions (100% ¹⁴N, 50% ¹⁴N and 50% ¹⁵N, and 100% ¹⁵N) for 6 weeks. Regardless of treatment, all available inorganic N was taken up by M. spicatum at similar rates. Significant differences in tissue isotopic composition were evident across treatments in 2 weeks, and persisted for the remainder of the experiment. At 6 weeks, atom (at) % ¹⁵N was twice as high in the 100% ¹⁵N treatment as in the 50% ¹⁵N treatment, and the at% of the ¹⁴N treatment (0% ¹⁵N) remained unchanged. This observed absence of preference for ¹⁴N in M. spicatum suggests that this species can be useful in determining N sources to freshwater systems, which often differ in ¹⁵N/¹⁴N ratio.     

Sensitivity of the Green Alga Pediastrum duplex Meyen to Allelochemicals Is Strain-Specific and Not Related to Co-Occurrence with Allelopathic Macrophytes

Interspecific differences in the response of microalgae to stress have numerous ecological implications. However, little is known of intraspecific sensitivities and the potential role of local genetic adaptation of populations. We compared the allelochemical sensitivity of 23 Pediastrum duplex Meyen strains, a common component of the freshwater phytoplankton. In order to test for local genetic adaptation, strains were isolated from water bodies with and without the allelopathically-active submerged macrophyte Myriophyllum. Strains were assigned to P. duplex on the basis of cell shape and colony morphology and only P. duplex strains that belonged to the same lineage in an ITS rDNA phylogeny were used. Inhibition of strain growth rates and maximum quantum yields of photosystem II were measured after exposure to tannic acid (TA) and co-culture with Myriophyllum spicatum. Growth rate inhibition varied over one order of magnitude between the P. duplex strains. There was no correlation between the presence of Myriophyllum in the source location and the sensitivity of the strains to TA or the presence of Myriophyllum, suggesting that at least strong unidirectional local adaptation to Myriophyllum had not taken place in the studied water bodies. The maximum quantum yield of photosystem II of TA exposed algae decreased, whereas the yield of algae exposed to M. spicatum was slightly higher than that of the controls. The ranking of P. duplex strain sensitivities differed between the types of exposure (single additions of TA versus co-existence with M. spicatum) and the parameter measured (growth rate versus maximum quantum yield), emphasizing the importance of measuring multiple traits when analysing strain-specific sensitivities towards allelochemicals. The observation that sensitivities to allelochemicals vary widely among strains of a single freshwater algal species should be taken into account if evaluating ecological consequences of allelopathic interactions.     

Trisetum spicatum (L.)Richt. in northern Borneo

A new subspecies ofTrisetum spicatum (L.)Richt. has been described from Borneo.     

Phytoplankton as a factor in the decline of the submersed macrophyte Myriophyllum spicatum L. in Lake Wingra,Wisconsin, U.S.A.

A dramatic decline in biomass and areal coverage of the submersed macrophyte Myriophyllum spicatum in Lake Wingra, Wisconsin, USA during the mid-1970's is documented using aerial photography, vegetation surveys, and quadrat biomass sampling. Over the same period, light penetration as measured by Secchi disc transparency and extinction coefficient decreased substantially. During this period, extinction coefficient was closely correlated with chlorophyll a levels implicating phytoplankton as the major source of decreased light penetration. A growth model for M. spicatum predicts a substantial decline in macrophyte biomass when extinction coefficient is increased to the levels reported since 1977. Available data do not show whether the phytoplankton increase preceeded or followed the macrophyte decline. Nonetheless it is clear that phytoplankton growth can account for a substantial portion of the decline and that macrophyte recovery will be difficult given the phytoplankton-induced decrease in water clarity.     

The distribution ofMyriophyllum spicatum L. in the inland waters of Egypt

Myriophyllum spicatum has been discovered in the Nile valley during the past decade. The rapid colonization seems to be influenced by changes in the Nile water regime after the building of the High Dam at Aswan.