Effects of an exotic invasive macrophyte (tropical signalgrass) on native plant community composition,species richness and functional diversity

1. The issue of freshwater species being threatened by invasion has become central in conservation biology because inland waters exhibit the highest species richness per unit area, but apparently have the highest extinctions rates on the planet. 2. In this article, we evaluated the effects of an exotic, invasive aquatic grass (Urochloa subquadripara– tropical signalgrass) on the diversity and assemblage composition of native macrophytes in four Neotropical water bodies (two reservoirs and two lakes). Species cover was assessed in quadrats, and plant biomass was measured in further quadrats, located in sites where tropical signalgrass dominated (D quadrats) and sites where it was not dominant or entirely absent (ND quadrats). The effects of tropical signalgrass on macrophyte species richness, Shannon diversity and number of macrophyte life forms (a surrogate of functional richness) were assessed through regressions, and composition was assessed with a DCA. The effects of tropical signalgrass biomass on the likelihood of occurrence of specific macrophyte life forms were assessed through logistic regression. 3. Tropical signalgrass had a negative effect on macrophyte richness and Shannon and functional diversity, and also influenced assemblage composition. Emergent, rooted with floating stems and rooted submersed species were negatively affected by tropical signalgrass, while the occurrence of free‐floating species was positively affected. 4. Our results suggest that competition with emergent species and reduction of underwater radiation, which reduces the number of submersed species, counteract facilitation of free‐floating species, contributing to a decrease in plant diversity. In addition, homogenisation of plant assemblages shows that tropical signalgrass reduces the beta diversity in the macrophyte community. 5. Although our results were obtained at fine spatial scales, they are cause for concern because macrophytes are an important part of freshwater diversity.     

Effects of the density of the invasive macrophyte <Emphasis Type="Italic">Hydrilla verticillata</Emphasis> and root competition on growth of one native macrophyte in different sediment fertilities

The occurrence of non-native species at high densities may generate competition for resources and possibly exclude native species in various environments. We evaluated the effects of increased densities of the non-native invasive macrophyte Hydrilla verticillata on the growth of the native species Egeria najas in different sediment types and with only root interactions or root?+?shoot interactions. We tested the hypothesis that the effect of the invasive on the native species is density dependent and that it is greater when competition for light and nutrients occurs (root?+?shoot interactions). The results of these experiments demonstrated that increased density of the invasive species H. verticillata significantly decreased the growth of the native species independent of sediment type (sand or mud sediments). When plants competed for water and sediment resources (root?+?shoot interactions), the native species was more impacted by the invasive than when they competed only for water resources (only shoots interacting). Our results show that E. najas is probably unable to colonize sites highly colonized by hydrilla, and this applies to both sand and mud sediments. This outcome suggests that H. verticillata is a threat for E. najas and likely other native submerged species in South America.     

Differential influence of a monotypic and diverse native aquatic plant bed on a macroinvertebrate assemblage; an experimental implication of exotic plant induced habitat

Aquatic plants mediate ecological processes in aquatic habitats, specifically predator–prey (bluegill sunfish (Lepomis macrochirus Rafinesque)-macroinvertebrate) interactions. Macroinvertebrate colonization is directly and indirectly influenced by substrate heterogeneity, interstitial space, and surface complexity. Exotic invasive plant species, such as Hydrilla verticillata L.F. Royle, may alter the available structure in aquatic habitat by creating a shift to a homogeneous habitat, thus affecting the macroinvertebrate community. Since macroinvertebrates provide a food base for young phytophilic fishes, changes in their density and abundance may alter food webs. We investigated the hypothesis that macroinvertebrate community structure is influenced by differences in habitat heterogeneity by measuring difference between a heterogeneous native aquatic plant bed, homogenous hydrilla plant bed, and habitat with no plants. Studies were conducted in the field (pond) and the experimental treatments were: (1) no plants, (2) monotypic bed of hydrilla, and (3) diverse native plants. Aquatic plants, regardless of species, supported greater macroinvertebrate abundance, richness, and biomass. Macroinvertebrate abundance, richness, and biomass in a hydrilla-dominated habitat did not differ significantly from a diverse plant habitat, except for richness in October. Indicator taxa did differ significantly between respective treatments, suggesting a change in species composition. However, no significant effect of fish predation on macroinvertebrate populations and/or community structure was documented. The data suggest that a shift from a natural mosaic of vegetated habitat to a highly complex monotypic habitat (e.g., exotic hydrilla) may reduce spatial heterogeneity important to structuring a macroinvertebrate assemblage. Handling editor: S. M. Thomaz     

Aquatic macrophytes in the large, sub-tropical Itaipu Reservoir, Brazil

In the last three decades, rapid assessment surveys have become an important approach for measuring aquatic ecosystem biodiversity. These methods can be used to detect anthropogenic impacts and recognize local or global species extinctions. We present a floristic survey of the aquatic macrophytes along the Brazilian margin of the Itaipu Reservoir conducted in 2008 and compare this with a floristic survey conducted ten years earlier. We used ordination analysis to determine whether assemblage composition differed among reservoir arms. Macrophyte species were sampled in each of the 235 sampling stations using a boat, which was positioned inside three places of each macrophyte stand to record species and search for small plants. We also collected submerged plants using a rake with the boat moving at constant velocity for ten minutes. We assigned individual macrophyte species to life form and identified representative species for each life form. A total of 87 macrophyte taxa were identified. The "emergent" life forms contained the highest number of species, followed by "rooted submerged" life forms. The extensive survey of macrophytes undertaken in September 2008 recorded more species than a survey conducted between 1995 and 1998. This could be due to changes in water physico-chemistry, disturbances due to water drawdown and the long period between surveys, which may have allowed natural colonization by other species. Additionally, differences in the classification systems and taxonomic resolution used in the surveys may account for differences in the number of species recorded. Assemblage composition varied among the arms and was affected by underwater radiation (as measured using a Secchi disk) and fetch. Five non-native species were found. Two of these non-native species (Urochloa subquadripara and Hydrilla verticillata) are of special concern because they have a high frequency of occurrence and occupy large marginal areas of the reservoir. Future surveys should be conducted to determine the habitat most frequently colonized by these species. This would allow management strategies to be developed to protect native aquatic biota and prevent interference with the recreational and commercial uses of the Itaipu Reservoir.     

Interactions between bacteria and fungi in macrophyte leaf litter decomposition

Microbes play an important role in decomposition of macrophytes in shallow lakes, and the process can be greatly affected by bacteria–fungi interactions in response to material composition and environmental conditions. In this study, microbes involved in the decomposition of leaf litter from three macrophyte species, Zizania latifolia, Hydrilla verticillata and Nymphoides peltata, were analysed at temperatures of 5, 15 and 25 °C. Results indicate that the decomposition rate was affected by temperature. Bacterial alpha diversity increased significantly along the time, while both temperature and plant species had a significant impact on the bacterial community, and plant type was shown to be the most important driving factor for the fungal community. The cosmopolitan bacterial taxa affiliated with Gammaproteobacteria, Bacteroidetes, Deltaproteobacteria, Firmicutes and Spirochaetes were key species in the investigated ecological networks, demonstrating significant co-occurrence or co-exclusion relationships with Basidiomycota and Ascomycota, according to different macrophyte species. This study indicates that bacteria involved in the decomposition of macrophyte leaf litter are more sensitive to temperature variance, and that fungi have a higher specificity to the composition of plant materials. The nutrient content of Hydrilla verticillata promoted a positive bacteria–fungi interaction, thereby accelerating the decomposition and re-circulation of leaf litter.     

Effects of desiccation on the toxicant sensitivity of rotifers

Non-native aquatic macrophytes have invaded different types of ecosystems all over the world. The exotic submersed macrophyte Hydrilla verticillata recently invaded the Paraná basin, Brazil, being recorded by the first time in the natural habitats of this river in 2005. We investigated the effects of this species on ostracod assemblages and compared the abundance, richness, and Shannon–Wiener diversity of ostracod assemblages that colonize the invading species with those that colonize Egeria najas, a native submersed species with similar architecture and physical complexity. Fragments of these two species were left for 28 days in tanks to root and grow and then they were transferred to a floodplain lake where they remained in pairs (one plant of each species; N = 7) during 30 days for colonization by ostracods. A detrented correspondence analysis was used to summarize ostracod assemblage composition. Although there were no significant differences in ostracod abundance, richness and Shannon diversity when analyzed separately, cumulative curves, which permit to eliminate effects of abundance on richness, indicated a significantly higher number of ostracod species on H. verticillata. Assemblage composition was significantly different between both plant species, as shown by the first DCA axis. Our results show that H. verticillata might provide favorable habitats for native ostracod assemblages.     

Do Amplitudes of Water Level Fluctuations Affect the Growth and Community Structure of Submerged Macrophytes?

Submerged macrophytes are subjected to potential mechanical stresses associated with fluctuating water levels in natural conditions. However, few experimental studies have been conducted to further understand the effects of water level fluctuating amplitude on submerged macrophyte species and their assemblages or communities. We designed a controlled experiment to investigate the responses of three submerged macrophyte species (Hydrilla verticillata, Ceratophyllum demersum and Elodea nuttallii) and their combinations in communities to three amplitudes (static, ± 30 cm, ± 60 cm) of water level fluctuations. Results showed that water level fluctuating amplitude had little effects on the community performance and the three tested species responded differently. H. verticillata exhibited more growth in static water and it was negatively affected by either of the water level fluctuations amplitude, however, growth parameters of H. verticillata in two fluctuating water level treatments (i.e., ± 30 cm, ± 60 cm) were not significantly different. On the other hand, the growth of C. demersum was not significantly correlated with different amplitude treatments. However, it became more abundant when water levels fluctuated. E. nuttallii was inhibited by the two fluctuating water level treatments, and was less in growth parameters compared to the other species especially in water level fluctuating conditions. The inherent differences in the adaptive capabilities of the tested species indicate that C. demersum or other species with similar responses may be dominant species to restore submerged macrophyte communities with great fluctuating water levels. Otherwise, H. verticillata, E. nuttallii or other species with similar responses could be considered for constructing the community in static water conditions.     

High invasion potential of Hydrilla verticillata in the Americas predicted using ecological niche modeling combined with genetic data

Ecological niche modeling is an effective tool to characterize the spatial distribution of suitable areas for species, and it is especially useful for predicting the potential distribution of invasive species. The widespread submerged plant Hydrilla verticillata (hydrilla) has an obvious phylogeographical pattern: Four genetic lineages occupy distinct regions in native range, and only one lineage invades the Americas. Here, we aimed to evaluate climatic niche conservatism of hydrilla in North America at the intraspecific level and explore its invasion potential in the Americas by comparing climatic niches in a phylogenetic context. Niche shift was found in the invasion process of hydrilla in North America, which is probably mainly attributed to high levels of somatic mutation. Dramatic changes in range expansion in the Americas were predicted in the situation of all four genetic lineages invading the Americas or future climatic changes, especially in South America; this suggests that there is a high invasion potential of hydrilla in the Americas. Our findings provide useful information for the management of hydrilla in the Americas and give an example of exploring intraspecific climatic niche to better understand species invasion.     

HYDRILLA INVADES WASHINGTON,D.C. AND THE POTOMAC

The aquatic weed hydrilla (Hydrilla verticillata Royle) was discovered growing in the Potomac River, south of Alexandria, VA, in Kenilworth Aquatic Gardens, Washington, D.C., and in the Chesapeake and Ohio (C&O) Canal near Seneca, MD. Cultures in Florida of the Kenilworth clone produced male flowers. This is the first report of the occurrence of the male in the U.S. Two distinct isoenzyme patterns have been identified for plants from various locations in the U.S., corresponding to a monoecious strain and a dioecious female. The occurrence of the wild colonies of the monoecious Hydrilla greatly increase the potential for physiological diversity through sexual reproduction, which may have serious consequences for the management of this weed.     

Using space‐for‐time substitution and time sequence approaches in invasion ecology

1. Invasion biologists use two main approaches to evaluate the effects of non‐native species (NNS) on diversity of native species (DNS), namely space‐for‐time and time approaches. These approaches have pitfalls related to lack of controls: the former lacks pre‐invasion data, while the latter often lacks data from non‐invaded sites. 2. We propose a framework that combines space‐for‐time and time approaches and which should result in more focused mechanistic hypotheses and experiments to test the causes of invasibility and the effects of NNS on DNS. We illustrate the usefulness of our framework using two case studies: one with the submersed macrophyte, Hydrilla verticillata, in reservoir and the other with the fish, Geophagus proximus, in a large river–floodplain system. 3. Hydrilla verticillata invaded sites with DNS similar to that found in non‐invaded sites, indicating that biotic and/or abiotic factors did not influence invasion success; however, DNS increased over time in invaded sites compared with non‐invaded sites, suggesting that H. verticillata facilitated natives. In contrast, G. proximus invaded sites with higher DNS than non‐invaded sites, suggesting that biotic and/or abiotic factors favouring natives were important for invasion success, but DNS increased in invaded and non‐invaded sites over time, indicating that an independent factor contributed to DNS increases. 4. Conclusions from both studies would have been inaccurate or incomplete if the space‐for‐time and time approaches had not been used in combination as proposed in our framework.     

Environmental and Chemical Control of Vegetative Dormant Bud Production in Hydrilla verticillata

Bud production was investigated using algal-free cultures ofthe aquatic weed, Hydrilla verticillata. Vegetative dormantbud production in hydrilla is stimulated by a shortened photoperiodand is not temperature-dependent between 15 and 30 C. The phytochromesystem appears to be involved in this process and ABA stimulatesbud production. Artificial illumination throughout the nightprevented dormant bud production. Ethylene, applied as ethephon,reduced bud production in greenhouse cultures by 80 per centwhen applied at a level of 0.1 mgl^–1 ethephon at 2-d intervals. Hydrilla verticillata, aquatic plants, dormant buds, ethephon, ethylene, abscisic acid     

Impacts of sediment type on the performance and composition of submerged macrophyte communities

To restore deteriorated lake ecosystems, it is important to identify environmental factors that influence submerged macrophyte communities. While sediment is a critical environmental factor for submerged macrophytes and many studies have examined effects of sediment type on the growth of individual submerged macrophytes, very few have tested how sediment type affects the growth and species composition of submerged macrophyte communities. We constructed submerged macrophyte communities containing four co-occurring submerged macrophytes (Hydrilla verticillata, Myriophyllum spicatum, Ceratophyllum demersum and Chara fragilis) and subjected them to three sediment treatments, i.e., clay, a mixture of clay and quartz sand at a volume ratio of 1:1 and a mixture at a volume ratio of 1:4. Compared to the clay, the 1:1 mixture treatment greatly increased overall biomass, number of shoot nodes and shoot length of the community, but decreased its diversity. This was because it substantially promoted the growth of H. verticillata within the community, making it the most abundant species in the mixture sediment, but decreased that of M. spicatum and C. demersum. The sediment type had no significant effects on the growth of C. fragilis. As a primary nutrient source for plant growth, sediment type can have differential effects on various submerged macrophyte species and 1:1 mixture treatment could enhance the performance of the communities, increasing the overall biomass, number of shoot nodes and shoot length by 39.03%, 150.13% and 9.94%, respectively, compared to the clay treatment. Thus, measures should be taken to mediate the sediment condition to restore submerged macrophyte communities with different dominant species.     

Behavioural response of fishes to increasing pH and dissolved oxygen: field and laboratory observations

• 1 Freshwater fish assemblages were sampled quantitatively before, during, and after naturally produced conditions of elevated pH and dissolved oxygen (DO) that occurred for 2–6h in two dense macrophyte beds. Field observations were followed by 3–4-h laboratory behavioural experiments in which banded killifish (Fundulus diaphanus), bluegill (Lepomis macrochirus), and striped bass (Morone saxatilis) were exposed to gradually increasing levels of: (i) pH; (ii) DO; and (iii) pH and DO.
• 2 Field results indicated that the macrophyte bed dominated by Hydrilla verticillata had a higher fish density than the mixed-species (H. verticillata and Vailisneria americana) bed, but beds did not differ significantly in fish species richness or biomass. Within beds, diel changes in fish species composition, richness, density and biomass were minor.
• 3 In the laboratory, avoidance of pH levels exceeding 9.5 was exhibited by all species tested. No significant response was observed to either DO increase treatments, or to pH and DO increase treatments.
• 4 Results suggest fishes do not avoid areas, such as dense macrophyte beds, when high pH levels (9.5–10.0) are accompanied by high DO levels (200–260% saturation).

     

Effects of hydrological variation on the aquatic plant community in a floodplain palustrine wetland of southern Brazil

This study analyzed macrophyte richness, biomass, and composition under flooding of brief duration (less than 3 days) and drawdown events over an annual cycle in a floodplain palustrine wetland in the south of Brazil. The study was carried out to test the hypothesis that floods of brief and very brief duration are not long enough to compromise the richness and the biomass of aquatic macrophytes and that the alternation between wet and drawdown phases may cause variations in the macrophyte richness and composition. A total of 26 aquatic macrophyte species were observed from April 2003 to May 2004: 13 species were observed during the wet phase, and 24 during the drawdown phase. The mean richness was higher during the drawdown phase than during the wet phase, however, the mean biomass was similar in both phases. Although macrophyte richness was not modified after the three flooding events, mean biomass was modified after two events. The number of macrophyte species of which the biomass was modified after the first flooding event increased with subsequent floods. These results illustrate the importance of the dynamics between brief floods and drawdown events to the aquatic plant community in floodplain wetlands in southern Brazil.     

Host range and searching behaviour of Cricotopus lebetis (Diptera: Chironomidae), a tip miner of Hydrilla verticillata (Hydrocharitaceae)

A chironomid midge, Cricotopus lebetis Sublette (Diptera: Chironomidae), was discovered feeding on Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) in Crystal River, Citrus, Co., Florida, in the 1990s. Larvae of the midge mine the apical meristems of hydrilla, causing terminal branching and stunting of the plant. We investigated the fundamental host range of the midge by conducting a series of no-choice and paired-choice tests. No-choice developmental tests with neonate larvae revealed that the fundamental host range of C. lebetis included not only on hydrilla but also several other aquatic plants in different families, suggesting that this insect is not a hydrilla specialist. In paired-choice bioassays, larval colonisation of Elodea canadensis Michx. (Hydrocharitaceae) and Najas guadalupensis (Spreng.) Magnus (Najadaceae) was greater than colonisation of H. verticillata. Behavioural bioassays in a Y-tube olfactometer and in Petri dishes suggested that neonate larvae were not able to locate host plant material, whereas older larvae were successful in finding hosts. In paired-choice oviposition tests, adult females discriminated between potential oviposition sites, with greater numbers of eggs laid on E. canadensis and N. guadalupensis than on H. verticillata. This study is the first detailed account of host searching and oviposition behaviour of a phytophagous chironomid midge. The results will be used to assess the potential value of C. lebetis as a biological control agent of hydrilla.     

Competitive Interactions between Hydrilla (Hydrilla verticillata) and Vallisneria (Vallisneria americana) as Influenced by Insect Herbivory

Two experiments (winter and summer) were conducted in outdoor tanks using addition-series methods to evaluate the impact of specialized feeding by two biological control agents,Hydrellia pakistanaeDeonier andBagous hydrillaeO'Brien, on competitive interactions between hydrilla [Hydrilla verticillata(L.f.) Royle] and vallisneria (Vallisneria americanaMichx). Competitive abilities of each plant species were determined using the reciprocal-yield model of mean plant weight. In the absence of the biocontrol agents, intraspecific competition from hydrilla on itself was 8.3 times stronger than interspecific competition from vallisneria.Hydrellia pakistanaeinterfered with hydrilla canopy formation by removing as much as 80% of the plant biomass in the top 30 cm of the water column. Damage byH. pakistanaealso caused a 43% reduction in hydrilla tuber production during the winter experiment. Similarly,B. hydrillaecaused up to a 48% reduction in hydrilla plant weight in the summer experiment. Neither insect species damaged vallisneria. As a result, there were significant shifts in the competitive balance between hydrilla and vallisneria due to selective insect feedings. In the presence ofH. pakistanae, hydrilla intraspecific competition was nearly equal to interspecific competition from vallisneria, indicating that hydrilla had lost its competitive edge over vallisneria.Bagous hydrillaealso produced similar, but smaller, shifts in the relative competitive abilities of hydrilla and vallisneria. These results indicate that biological control agents can disrupt the competitive balance between plant species in favor of native species, thus adding another element to the weed biological control strategies.     

Geographic selection bias of occurrence data influences transferability of invasive Hydrilla verticillata distribution models

Due to socioeconomic differences, the accuracy and extent of reporting on the occurrence of native species differs among countries, which can impact the performance of species distribution models. We assessed the importance of geographical biases in occurrence data on model performance using Hydrilla verticillata as a case study. We used Maxent to predict potential North American distribution of the aquatic invasive macrophyte based upon training data from its native range. We produced a model using all available native range occurrence data, then explored the change in model performance produced by omitting subsets of training data based on political boundaries. We also compared those results with models trained on data from which a random sample of occurrence data was omitted from across the native range. Although most models accurately predicted the occurrence of H. verticillata in North America (AUC > 0.7600), data omissions influenced model predictions. Omitting data based on political boundaries resulted in larger shifts in model accuracy than omitting randomly selected occurrence data. For well‐documented species like H. verticillata, missing records from single countries or ecoregions may minimally influence model predictions, but for species with fewer documented occurrences or poorly understood ranges, geographic biases could misguide predictions. Regardless of focal species, we recommend that future species distribution modeling efforts begin with a reflection on potential spatial biases of available occurrence data. Improved biodiversity surveillance and reporting will provide benefit not only in invaded ranges but also within under‐reported and unexplored native ranges.     

Response of native Egeria najas Planch. and invasive Hydrilla verticillata (L.f.) Royle to altered hydroecological regime in a subtropical river

Egeria najas Planch. is the dominant native submersed macrophyte of the Upper Paraná River in Brazil, while Hydrilla verticillata (L.f.) Royle has recently invaded this area. From January 2006 to December 2007, comprising two annual flood cycles, we conducted monthly surveys at two river stations and two lakes connected to the river within this stretch of the Paraná River, aiming to understand how the hydrological regime influences the distribution and abundance of these native and invasive Hydrocharitaceae species. Hydrilla did not develop in the lakes, possibly due to the elevated proportion of organic matter in the sediment (∼10% DW). However, the exotic species dominated the river sites apparently suppressing E. najas. In the lakes E. najas reached a maximum biomass of 628 ± 82 g DW m⁻² but did not surpass 333 ± 83 g DW m⁻² in the river, where H. verticillata peaked at 1415 ± 255 g DW m⁻². Macrophyte biomass development was greatest during low-water periods, with transparent water and high temperatures. Floods probably affected submersed macrophytes (especially in 2007, when an extreme flood caused by an El Niño Southern Oscillation (ENSO) event occurred) via sediment movement and plant scouring (uprooting) effects, coupled with reduced water transparency. Macrophyte recovery started soon after the (less intense) 2006 flood but was delayed in 2007. In the river recovery started five months after the major flood, but in the lakes no significant plant regeneration was found even nine months after the disturbance. E. najas and H. verticillata started regeneration practically at the same time but H. verticillata had much higher rates of biomass increase.     

Response of invasive macrophyte species to drawdown: The case of Elodea sp.

The response to drawdown of vegetative fragments (whole plants, shoot fragments and turions) of two invasive macrophyte species, Elodea canadensis and Elodea nuttallii, was studied through laboratory experiments. In addition, field observations were made on the colonisation of a wetland by E. nuttallii before and after a natural drawdown. The survival and the growth of vegetative fragments of E. nuttallii were higher than those of E. canadensis after an artificial drawdown of several days. In the field the recolonisation by E. nuttallii of a wetland that was drained for 10 weeks during a summer drawdown was very rapid, the abundance of this macrophyte species being not affected by the drawdown event. We conclude that E. nuttallii possesses a high resilience to desiccation and that a summer drawdown would not be efficient in the control of this invasive species.     

Response of invasive macrophyte species to drawdown: The case of Elodea sp.

The response to drawdown of vegetative fragments (whole plants, shoot fragments and turions) of two invasive macrophyte species, Elodea canadensis and Elodea nuttallii, was studied through laboratory experiments. In addition, field observations were made on the colonisation of a wetland by E. nuttallii before and after a natural drawdown. The survival and the growth of vegetative fragments of E. nuttallii were higher than those of E. canadensis after an artificial drawdown of several days. In the field the recolonisation by E. nuttallii of a wetland that was drained for 10 weeks during a summer drawdown was very rapid, the abundance of this macrophyte species being not affected by the drawdown event. We conclude that E. nuttallii possesses a high resilience to desiccation and that a summer drawdown would not be efficient in the control of this invasive species.