Limits and effects of invasion by the nonindigenous wetland plant <Emphasis Type="Italic">Lythrum salicaria</Emphasis> (purple loosestrife): a seed bank analysis

We used seed bank analyses to investigate the role of dispersal in limiting invasion by Eurasian Lythrum salicaria within and among North American wetlands, and the changes in seed bank diversity associated with this invader. We compared the number and species composition of seedlings emerging from soil sampled in 11 uninvaded wetlands and paired uninvaded and invaded sites within 10 invaded wetlands under both seedling competition and noncompetitive conditions. Almost no L. salicaria emerged in samples from uninvaded wetlands, indicating dispersal limitation despite prodigious seed production in nearby wetlands. However L. salicaria emerged in all samples from uninvaded sites in invaded wetlands, suggesting environmental limits on establishment within invaded wetlands. Conditions that provided opportunities for seedlings to compete reduced survival of Typha spp. but not L. salicaria seedlings. However, this was due to species-specific differences in post-emergence mortality rather than response to competition. Competition did reduce seedling mass, but this effect did not differ among species. Species richness of emerging seedlings was lower for invaded than uninvaded wetlands. Lower seed bank richness may be a cause or consequence of L. salicaria invasion. Efforts to reduce seed dispersal to uninvaded wetlands would likely slow the spread of this invader.     

Impacts of major predators on tropical agroforest arthropods: comparisons within and across taxa

Non-native plants can have adverse effects on ecosystem structure and processes by invading and out-competing native plants. I examined the hypothesis that mature plants of non-native and native species exert differential effects on the growth of conspecific and heterospecific seedlings by testing predictions that (1) invasive vegetation has a stronger suppressive effect on seedlings than does native vegetation, (2) seedlings of invasive species are better able to grow in established vegetation than are native seedlings, and (3) invasive species facilitate conspecific and inhibit heterospecific seedling growth. I measured growth rates and interaction intensities for seedlings of four species that were transplanted into five wetland monoculture types: invasive Lythrum salicaria; native L. alatum, Typha angustifolia, T. latifolia; unvegetated control. Invasive L. salicaria had the strongest suppressive effect on actual and per-individual bases, but not on a per-gram basis. Seedlings of T. latifolia were better able to grow in established vegetation than were those of L. salicaria and T. angustifolia. These results suggest that L. salicaria is not a good invader of established vegetation, but once established, it is fairly resistant to invasion. Thus, it is likely that disturbance of established vegetation facilitates invasion by L. salicaria, allowing it to compete with other species in even-aged stands where its high growth rate and consequent production of aboveground biomass confer a competitive advantage.     

Decomposition rates and phosphorus concentrations of purple Loosestrife (Lythrum salicaria) and Cattail (Typha spp.) in fourteen Minnesota wetlands

Purple Loosestrife is rapidly displacing native vegetation in North American wetlands. Associated changes in wetland plant communities are well understood. Effects of Loosestrife invasion on nutrient cycling and decomposition rates in affected wetlands are unknown, though potentially of significance to wetland function. We used litter bag methods to quantify decomposition rates and phosphorus concentrations of purple Loosestrife (Lythrum salicaria) and native cattails (Typha spp.) in fourteen Minnesota wetlands. A 170-day study that began in autumn modeled decomposition of Loosestrife leaves. Loosestrife stems andTypha shoots that had overwintered and fragmented were measured in a 280- day study that began in spring. In general, Loosestrife leaves decomposed most rapidly of the three;Typha shoots decomposed faster than Loosestrife stems. Significant decay coefficients (k-values) were determined by F-testing single exponential model regressions of different vegetation types in the fourteen wetlands. Significant decay coefficients were:k = 2.5 × 10⁻³ and 4.32 × 10⁻³ for all Loosestrife leaves (170 d);k = 7.2 × 10⁻⁴ and 1.11 × 10⁻³ for overwintered Loosestrife stems (280-d) andk = 7.9 × 10⁻⁴, 1.42 × 10⁻³ and 2.24 × 10⁻³ for overwinteredTypha shoots (280-d). Phosphorus concentrations of plant tissue showed an initial leaching followed by stabilization or increase probably associated with microbial growth. Loosestrife leaves had twice the phosphorus concentration of Loosestrife stems andTypha shoots. Our results indicate that conversion of wetland vegetation from cattails to Loosestrife may result in significant change in wetland function by altering timing of litter input and downstream phosphorus loads. Conversion of a riverine, flow- through wetland fromTypha to Loosestrife may effectively accelerate eutrophication of downstream water bodies. Impacts of Loosestrife invasion must be considered when wetlands are managed for wildlife or for improvement of downstream water quality.     

Constraints on Sedge Meadow Self‐Restoration in Urban Wetlands

Invasive plants and urban run‐off constrain efforts to restore sedge meadow wetlands. We asked if native graminoids can self‐restore following the removal of Typha × glauca (hybrid cattail), and if not, what limits their recovery? After we harvested Typha and depleted its rhizome starch reserves, Carex spp. expanded vegetatively (approximately 1 m over 2 years) but not by recruiting seedlings. A seedling emergence experiment showed that seed banks were depleted where Typha had eliminated the sedge meadow over a decade ago (based on aerial photo analysis). Carex seedling emergence was 75–90% lower where Carex was absent than where it remained in the plant community, and at least 17 species that were abundant 30 years ago were absent from the seed bank and extant vegetation. By varying hydroperiod, we showed that prolonged flooding prevented emergence of Carex seedlings and that a fluctuating hydroperiod reduced emergence and ultimately killed all Carex seedlings. In contrast, Typha seedlings emerged and survived regardless of hydroperiod. Thus, slow vegetative expansion by Carex, depauperate seed banks, and altered hydroperiods all constrain self‐restoration. To compensate for multiple constraints on self‐restoration, we recommend a long‐term management approach that capitalizes on flooding and the capacity of Carex spp. to regrow vegetatively. We suggest annually harvesting swaths of Typha at the edges of clones, before or during flood events, to allow gradual, vegetative self‐restoration of Carex spp.     

Mechanisms of dominance by the invasive hybrid cattail Typha?×?glauca

The mechanisms by which invasive plants displace native species are often not well elucidated, limiting knowledge of invasion dynamics and the scientific basis for management responses. Typha × glauca Godr. invades wetlands throughout much of North America. Like other problematic wetland invaders, Typha is large, grows densely, and leaves behind copious litter. It thus has the potential to impact wetlands both in life and after death. We assessed patterns in field settings and used simulated wetland-plant communities to experimentally test abiotic and community responses to live Typha, Typha litter, and water-level differences (confounded in the field) using a full-factorial design. In general, litter was a stronger driver of change than live Typha. The greatest impacts were seen where, as in nature, live and dead Typha co-occurred. Live-Typha treatments did not differ from controls in light or temperature conditions but did reduce community biomass and alter community composition. Litter strongly affected light, temperature and its variability, community and species-level plant biomass, and community composition. Interactions between live Typha and litter affected aspects of plant-community composition. Advantageously for Typha, interspecific litter effects were not mirrored by intraspecific suppression of live Typha. These findings clarify how Typha is such an effective invader. Similar mechanisms are likely involved in invasions by other plant species, particularly in wetlands. Managers should respond quickly to new Typha invasions and, when dealing with established stands, remove litter in addition to eradicating live plants.     

Effects of experimental harvesting of an invasive hybrid cattail on wetland structure and function

Invasive plants, such as the hybrid cattail Typha × glauca, can reduce biodiversity and alter the ability of wetlands to provide critical ecosystem services, including nutrient cycling and carbon storage. Several approaches have been used to reduce Typha dominance and restore invaded wetlands, but long‐term studies assessing benefits of these restoration efforts are limited. A previous study demonstrated that aboveground harvesting of Typha × glauca stems and litter reduced Typha dominance 2 years post‐treatment in a Great Lakes coastal wetland. In the current study, we extended monitoring of experimental aboveground Typha harvest to 4 years post‐treatment and added assessments of treatment effects on soil nutrients, carbon emissions, and microbial community composition. Aboveground harvest treatment resulted in a dramatic reduction in Typha litter cover that persisted for 4 years, increased soil temperature, and increased abundance of the native plant genus Carex. However, aboveground harvest treatment did not significantly reduce Typha abundance, nor did it have significant effects on soil nutrient concentrations, carbon fluxes, or the taxonomic composition of soil microbial communities. We did observe differences in bacterial community composition between plots based on time since Typha invasion, which may indicate some legacy effects of Typha invasion. At the scale of this experiment (4 × 4 m plots), our results indicate that a single aboveground removal of Typha × glauca is not sufficient to restore a heavily invaded freshwater wetland ecosystem, and that periodic harvesting of Typha stems and litter may be required to maintain native plant abundance.     

Reproductive strategy of an invasive thistle: effects of adults on seedling survival

Invasive species are known for their ability to form monocultures that exclude native species, yet intraspecific interactions among invasives have not been well studied. Cynara cardunculus (L.) is an invasive perennial thistle that establishes high-density populations in coastal California grasslands. We examined the natural distribution of C. cardunculus seedlings in an established population and found that nearly 100% of seedlings grew within 2 m of adults despite an expected distribution peak at 3 m from source plants based on measured dispersal distances. We then investigated the role of mature plants in seedling survival and establishment with regard to live vegetation, litter, and seedling distance by planting seedlings at increasing distances around adults and applying removal treatments to the focal adult rosettes. We applied control (no removal), adult rosette removal (live leaves), litter removal (dead leaves), and adult rosette plus litter removal (all aboveground plant material) treatments. Seedlings experienced a higher rate of survival, measured by senescence date, and establishment, measured by return rate the following year, with all adult rosette removal treatments. Inhibition by adult rosettes was reduced with distance to 60–80 cm from the rosette, and there was little effect of adult plants between 80 and 200 cm. These results suggest that adult rosettes may inhibit conspecific seedlings at very close distances but provide a favorable environment for seedlings within nearby interspaces. This pattern may contribute to the creation and maintenance of high-density populations in C. cardunculus. Land managers seeking to control this species may improve long-term effectiveness by expanding management efforts to include a 2 m radius around adult plants and treating within 5 months of seedling emergence to prevent recruitment rather than treating adults alone.     

Rehabilitation of Swamp Paperbark (<Emphasis Type="Italic">Melaleuca ericifolia</Emphasis>) wetlands in south-eastern Australia: effects of hydrology,microtopography, plant age and planting technique on the success of community-based revegetation trials

Wetlands dominated by Swamp Paperbarks (Melaleuca spp., Myrtaceae) are common in coastal regions across Australia. Many of these wetlands have been filled in for coastal development or otherwise degraded as a consequence of altered water regimes and increased salinity. Substantial resources, often involving community groups, are now being allocated to revegetating the remaining wetland sites, yet only rarely is the effectiveness of the rehabilitation strategies or on-ground procedures robustly assessed. As part of a larger project investigating the condition and rehabilitation of brackish-water wetlands of the Gippsland Lakes, we overlaid a scientifically informed experimental design on a set of community-based planting trials to test the effects of water depth, microtopography, plant age and planting method on the survival and growth of seedlings of Melaleuca ericifolia Sm. in Dowd Morass, a degraded, Ramsar-listed wetland in south-eastern Australia. Although previous laboratory and greenhouse studies have shown M. ericifolia seedlings to be salt tolerant, the strongly interactive effects of waterlogging and salinity resulted in high seedling mortality (>90%) in the field-based revegetation trials. Seedlings survived best if planted on naturally raised hummocks vegetated with Paspalum distichum L. (Gramineae), but their height was reduced compared with seedlings planted in shallowly flooded environments. Age of plants and depth of water were important factors in the survival and growth of M. ericifolia seedlings, whereas planting method seemed to have little effect on survival. Improved testing of revegetation methods and reporting of success or otherwise of revegetation trials will improve the effectiveness and accountability of projects aiming to rehabilitate degraded coastal wetlands.     

Invasive Plant and Experimental Venue Affect Tadpole Performance

Introductions of non-native predators and competitors appear to contribute to worldwide amphibian declines; however, potential negative impacts of invasive plants on habitat quality and amphibian populations have not been examined. Loss of diversity and alterations in ecosystem function associated with plant invasions may disrupt food webs, potentially leading to further declines of already threatened amphibian populations. We used a combination of small bins, mesocosms, and field experiments to examine the impacts of Eurasian purple loosestrife (Lythrum salicaria) replacing native cattails (Typha latifolia) in North American freshwater wetlands on survival, developmental rate, and diet (freshwater algae) of American toad (Bufo americanus) tadpoles. Tadpoles developed slower in L. salicaria compared to tadpoles developing in T. latifolia. This effect was consistent across experimental venues, although mesocosms showed this effect only in the second year of our study. Survival and development rates were always more variable in purple loosestrife than in cattail. In bins, tadpoles showed significantly reduced survival when raised in purple loosestrife extract and addition of leaf litter exacerbated this negative effect. Tadpole survival rates in mesocosms and field cages were not significantly different between plant species, most likely an effect of high variability among replicates. We suspect a combination of direct toxicity of high tannin concentrations in L. salicaria leaves and their indirect negative impacts on aquatic food webs are responsible for these results. Tadpole gut analyses revealed differences in algal communities among venues and between L. salicaria and T. latifolia suggesting that alterations in tadpole food quality and quantity contribute to the observed reduced tadpole performance. The replacement of native wetland plant species by L. salicaria does not represent a simple exchange of ecological equivalents and the function of invaded habitats for native species has clearly changed. While we were investigating only a single amphibian species, our results suggest that the impact of L. salicaria on ecosystem processes and aquatic food webs may be more general and likely to negatively affect other wetland species. The threats non-indigenous plants represent for amphibian populations and food webs may be underestimated, and warrant further investigation.     

Effect of experimental phosphorus enrichment on oligotrophic tropical marshes in Belize,Central America

Sedges from genus Eleocharisdominate extensive wetlands in the sugar cane growing areas of the Caribbean. Correlative data suggest that macrophytes in these wetlands are phosphorus limited. To determine effects of increased P input that can be expected, e.g. from agricultural runoff, a common sugar cane fertilizer was applied to representative plots in four marl-based and four peat-based marshes. The plots were located in the proximity of patches of Typha domingensis, which has been reported to be able to outcompete Eleocharis under nutrient rich conditions. Responses to the fertilizer treatment were documented as changes in: Plant height, density, biomass, net primary production, nutrient resorption, decomposition, plant and soil nutrient concentrations, percent cover of cyanobacterial mats, and potential colonization by Typha. Additions of phosphorus significantly increased plant density and height and, consequently, the aboveground net primary production. Phosphorus resorption efficiency following senescence was independent of fertilizer addition in Eleocharis but decreased in Typha from the fertilized plots. Phosphorus resorption proficiency was lower in fertilized plots for both Typha and Eleocharis. Decomposition of litter and cellulose assays was significantly faster in fertilized plots. No spontaneous establishment of Typha occurred in the fertilized plots, but survival of transplanted Typha was higher in fertilized plots than in controls. Increased plant density in fertilized plots led to elimination of a key component of these ecosystems, the nitrogen fixing cyanobacterial mats.     

Evaluation of carbamazepine uptake and metabolization by Typha spp., a plant with potential use in phytotreatment

Phytoremediation technologies such as constructed wetlands have shown higher efficiencies in removal of pharmaceuticals from wastewaters than conventional wastewater treatment processes, and plants seem to have an important role in the removal of some of those compounds. In this context, a study was conducted to assess tolerance, uptake, and metabolism of the epilepsy drug, carbamazepine, by the macrophyte Typha spp. This evaluation was conducted in hydroponic solutions with 0.5-2.0 mg/L of this pharmaceutical for a maximum period of 21 days. The removal of carbamazepine from nutrient solutions by the plants reached values of 82% of the initial contents. Furthermore, a metabolite (10,11-dihydro-10,11-epoxycarbamazepine) was detected in leaf tissues indicating carbamazepine translocation and metabolism inside plants. Activities of antioxidant enzymes catalase, superoxide dismutase, and guaiacol peroxidase generally increase (after some mild initial inhibition in the case of the latter enzyme) as result of the abiotic stress caused by the exposure to carbamazepine, but ultimately Typha seemed able to cope with its toxicity. The results obtained in this study suggest the ability of Typha spp., to actively participate in the removal of carbamazepine from water when used in phytotreatment systems.     

Invasive reed effects on benthic community structure in Lake Erie coastal marshes

We examined how dominance (% canopy cover) and invasion history of common reed, Phragmites australis, affected benthic macroinvertebrate diversity and density in 8 marshes along Lake Erie’s southern shoreline. We also compared macroinvertebrate densities among patches (0.25 m²) of reed, cattail (Typha spp.), and native flora (e.g., Sagittaria, Sparganium) and epiphytic algal communities on submerged stems of reed and cattail. Narrow-leaf cattail (T. angustifolia) is also a common invasive plant to these wetlands, but does not greatly change plant community composition or ecosystem conditions like reed. Macroinvertebrate diversity (Shannon–Weaver H′) was positively related to reed cover and was highest (4.6) in two marshes with ~35- and 5-year invasion histories. Shading from high reed cover increased H′-diversity, in part, by reducing the abundance of floating duckweed, which harbored many Hyalella azteca amphipods. Percent Ephemeroptera, Odonata, and Trichoptera was low to moderate across marshes, regardless of reed cover and invasion history. Macroinvertebrate density was not affected by reed cover or average plant stem density, and did not differ among plant types. However, epiphyton densities and % diatoms were greater on reed than on cattail, suggesting reed provides a better feeding habitat for microalgal grazers than Typha. Abundance rankings of common species in these diatom-dominated communities were also typically dissimilar between these plant types. Although % grazers was unrelated to epiphyton densities and % diatoms, grazer identity (snails) differed between natural and diked marshes, which had different microalgal food supplies. Our findings suggest that Phragmites does not necessarily adversely affect macroinvertebrate community structure and diversity and that invasion history alone has little effect on the H′-diversity–reed dominance relationship.     

Establishment of Carex stricta Lam. seedlings in experimental wetlands with implications for restoration

The loss of Carex dominated meadows due to agricultural drainage in the previously glaciated midcontinent of North America has been extensive. The lack of natural Carex recruitment in wetland restorations and the failures of revegetation attempts underscore the need for information on the establishment requirements of wetland sedges. In this study, seedlings of Carex stricta Lam. were planted in three experimental wetlands in east-central Minnesota, USA to investigate the biotic and abiotic environmental limitations to establishment. Seedlings were planted along an elevational water depth gradient to assess the effects of water depth and water level fluctuation on seedling survival and growth. A different water level fluctuation regime was assigned to each of the experimental wetlands to assess seedling tolerance for seasonal water level changes. The effects of seedling planting density and the presence or absence of non-sedge colonizers on seedling survival and growth were also studied. The experiment was followed for three growing seasons. The results of this study indicate that C. stricta seedlings were sensitive to the timing and duration of inundation during the first growing season. Once established, plants tolerated a broad range of seasonal drying and flooding conditions. Seedling and juvenile growth was slowed by non-sedge colonizers during the first two growing seasons, but by the third growing season, C. stricta was able to out-grow all annual and perennial weeds, except the aggressive perennial, Phalaris arundinacea L. The rapid growth of C. stricta plants, once established, indicates that the use of seedlings is a successful method for (re) introducing this tussock sedge into wetland restorations under a variety of environmental conditions. Comparison with other studies performed under similar conditions suggests that planting of seedlings is a more appropriate method of establishing this species than the use of transplanted rhizomes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Habitat and Population Effects on the Germination and Early Survival of the Invasive Weed, Lythrum salicaria L. (purple loosestrife)

In this study, we examined how environmental factors and differences among populations of Lythrum salicaria L. (Lythraceae; purple loosestrife) influenced the establishment of this species into both its preferred wetland habitat and secondary upland habitat. Seeds and seedlings from five parents from each of the three populations were planted in each wetland and upland of two sites in both disturbed and undisturbed plots. More seeds germinated in wetland than in upland sites (31% as compared to 11%) and, although a disturbance resulted in greater germination (23% as compared to 17% in undisturbed plots), there was no interaction with habitats. The results of a simultaneous greenhouse germination study in which seeds were planted under saturated and drained conditions suggest that the higher germination observed in wetland sites was due to the higher moisture present at such sites (79% vs. 45% in saturated and drained conditions, respectively). Disturbance enhanced both germination and growth of L. salicaria and this was especially so for seedlings. Dry weights of seedlings from disturbed plots were five times greater than those from undisturbed plots ( =9.2 vs. 1.8g in disturbed and undisturbed conditions, respectively). In addition to environmental factors, population differences are likely to play an important role in the spread of L. salicaria as we found differences between the populations in both life stages examined. One of the populations germinated equally well in both moisture treatments. In our seedling establishment experiment, one of the populations produced both the smallest plants and the most and earliest flowers, suggesting superior reproduction.     

Plant diversity in Lythrum,Phragmites, and Typhamarshes,Massachusetts, U.S.A.

Concern about colonization of marshesby plant species such as Phragmites australisand Lythrum salicariahas highlighted the needfor management strategies. However, there is a lack ofinformation in the literature on which to base thesedecisions. This study compares the alpha diversity ofmarshes to assess the impact of invasion by Phragmitesand Lythrum. Species occurrence andstem density were measured in marshes dominated by Phragmites, Lythrum, Typhaspp., or otherherbaceous perennials in the Charles River watershedin eastern Massachusetts, USA, and species richness,Shannon's H, Simpson's reciprocal (1/D), and Pielou'sJ were compared among six community types. The threediversity indices had significantly higher values forTypha-Lythrummarshes than for any of the othermarsh types (Tukey test, p< 0.05), with mean values(± s.d.) of H = 2.00 ± 0.74, 1/D = 3.51± 1.68 and J = 0.69 ± 0.1. Marshes dominatedby Phragmiteshad the lowest diversity, with H= 0 and D = 1, i.e. they were monospecific. Typhadominated marshes had the second lowest values,with H = 0.17 ± 0.05, 1/D = 1.05 ± 0.01, andJ = 0.11 ± 0.03. These results support the ideathat a reduction in diversity can be expected inmarshes colonized by Phragmites. However, thehigh diversity found in the Typha-Lythrummarshes contradicts the expectation of lower diversityafter invasion by Lythrum. This information mayalter marsh management decisions.     

Competitive effects of the invasive shrub,Lonicera maackii (Rupr.) Herder (Caprifoliaceae), on the growth and survival of native tree seedlings

Invasive plants are often associated with reduced cover of native plants, but rarely has competition between invasives and natives been assessed experimentally. The shrub Lonicera maackii, native to northeastern Asia, has invaded forests and old fields in numerous parts of eastern North America, and is associated with reduced tree seedling density in Ohio forests. A field experiment was conducted to test the effects of established L. maackii on the survival and growth of transplanted native tree species. The experiment examined above-ground competition (by removing L. maackii shoots) and below-ground competition (by trenching around transplanted seedlings). The effects of above-ground competition with L. maackii were generally more important than below-ground competition, though both were detected. Shoot treatment was the key determinant for the survival of all species except P. serotina, whereas trenching only enhanced survival for A. saccharum caged and P. serotina, and only in the shoot removal treatment. For the surviving seedlings, L. maackii shoot removal increased growth of A. saccharum seedlings protected with cages, but actually reduced the growth of unprotected Q. rubra and A. saccharum seedlings, indicating that L. maackii shoots confer some protection from deer browsing. Significant interactions between root and shoot treatment on Q. rubra growth parameters, specifically greatest growth in the shoot present & trenched treatment, is attributed to protection from deer browsing combined with release from below-ground competition. Despite this protective function of L. maackii shoots, the overall effect of this invasive shrub is increased mortality of native tree seedlings, suggesting it impacts the natural regeneration of secondary forests.     

Negative and positive interactions among plants: effects of competitors and litter on seedling emergence and growth of forest and grassland species

Living plant neighbours, but also their dead aboveground remains (i.e. litter), may individually exert negative or positive effects on plant recruitment. Although living plants and litter co‐occur in most ecosystems, few studies have addressed their combined effects, and conclusions are ambivalent. Therefore, we examined the response in terms of seedling emergence and growth of herbaceous grassland and forest species to different litter types and amounts and the presence of competitors. We conducted a pot experiment testing the effects of litter type (grass, oak), litter amount (low, medium, high) and interspecific competition (presence or absence of four Festuca arundinacea individuals) on seedling emergence and biomass of four congeneric pairs of hemicryptophytes from two habitat types (woodland, grassland). Interactions between litter and competition were weak. Litter presence increased competitor biomass. It also had positive effects on seedling emergence at low litter amounts and negative effects at high litter amounts, while competition had no effect on seedling emergence. Seedling biomass was negatively affected by the presence of competitors, and this effect was stronger in combination with high amounts of litter. Litter affected seedling emergence while competition determined the biomass of the emerged individuals, both affecting early stages of seedling recruitment. High litter accumulation also reduced seedling biomass, but this effect seemed to be additive to competitor effects. This suggests that live and dead plant mass can affect species recruitment in natural systems, but the mechanisms by which they operate and their timing differ.     

Yellow Bush Lupine Invasion in Northern California Coastal Dunes I. Ecological Impacts and Manual Restoration Techniques

We studied the ecological effects of the invasion of coastal dunes by Lupinus arboreus (yellow bush lupine), an introduced species, and used the results to develop manual restoration techniques on the North Spit of Humboldt Bay. Vegetation and soil data were collected in five vegetation types representing points along a continuum of bush lupine's invasive influence. We collected data on the number and size of shrubs, vegetation cover, and soil nutrients. One set of plots was subjected to two restoration treatments: removal of lupine shrubs only, or removal of all nonnative vegetation and removal of litter and duff. Treatments were repeated annually for four years, and emerging lupine seedlings were monitored for three years. Prior to treatment, ammonium and nitrate were found to increase along the lupine continuum, but organic matter decreased at the extreme lupine end. Yellow bush lupine was not the most significant variable affecting variation in soil nutrients. After four years, nonnative grasses, including Vulpia bromoides, Holcus lanatus (velvet grass), Bromus spp. (brome), and Aira spp. (European hairgrass), were significantly reduced in those restoration plots from which litter and duff was removed. Native species increased significantly in vegetation types that were less influenced by lupine. By the third year, soil variables differed among vegetation types but not by treatment. Bush lupine seedling emergence was higher, however, in plots receiving the litter and duff removal treatment. Based on these results, we conclude that bush lupine invasion results in both direct soil enrichment and indirect enrichment as a result of the associated encroachment of other nonnative species, particularly grasses. Although treatment did not affect soil nutrients during the period of this study, it did reduce establishment of nonnative grasses and recruitment of new bush lupine seedlings. Restoration should therefore include litter and duff removal. In areas that are heavily influenced by lupine and contain few native propagules, revegetation is also required.     

Litter Cover Variability Affects Seedling Performance and Herbivory1

We assessed the effects of litter cover on herbivory and performance (annual survival and relative growth rate in height and leaf number) for established seedlings (≥9 cm tall, and ≥4 mo old), of four shade-tolerant, large-seeded, co-occurring Sapotaceae species. Seedlings were divided into three litter treatments: (1) litter addition, (2) control, and (3) continuous litter removal. The mean survival for all treatments in descending order were: Chrysophyllum pomiferum (89%), Pouteria pemviemis (84%), P. caimito (75%), and Micropholis venulosa (40%). For all variables, species differed significantly within litter treatments. In M. venulosa, survival and relative growth rate in leaf number significantly decreased with the removal of litter, whereas herbivory increased. Relative growth rate in height of P. peruviensis decreased significantly with litter removal, while the levels of herbivory were higher with the addition of litter. Conversely, litter quantity had no effect on the performance of C. pomiferum, but herbivory was lower in the removal treatment. Finally, in P. caimito, litter treatment did not affect performance or herbivory. Results suggest that differences in litter quantity differentially affect tree species with similar biology (i.e., large-seeds, shade tolerance) in life stages other than germination and early establishment. The variability in species' responses to litter might be an important factor in determining species richness, abundance, and distribution of tropical rain forest tree species at the seedling level.     

Hydrologic alterations impact plant litter decay rate and ecosystem resilience in Mojave wetlands

Understanding ecosystem processes is vital for effective restoration of degraded ecosystems, especially wetlands. Restoration has become a necessity for management and conservation of the federally endangered Amargosa vole (Microtus californicus scirpensis) endemic to small, bulrush (Schoenoplectus americanus) dominated wetlands in the Mojave Desert. Recent data indicate catastrophic decrease of the vole population and its habitat from local alterations to hydrology, combined with diminished decomposition rates of bulrush, persistence of plant litter, and minimal plant growth except along narrow margins along stream edges. We conducted a series of three field and one greenhouse experiment(s) testing the effect of (1) moisture level on plant decay rate, (2) litter removal on plant regeneration, (3) the interactive effect of litter removal and moisture level increase on plant regeneration, and (4) potential germination rate of bulrush seeds under multiple hydrologic regimes to understand how hydrologic alteration and litter decay ultimately influences marsh regeneration. Results revealed decrease in water level caused a 20‐fold reduction in decomposition rates of a degraded marsh. Litter removal alone and in combination with water table restoration significantly and positively affected bulrush resprouting (p < 0.0001 for both). Seed bank experiments showed high rates of germination in saturated and flooded soil conditions, emphasizing the potential role of seedlings in ecosystem recovery. This study shows how the interaction of hydrologic change and decreased decomposition can shift an ecosystem toward limits of resilience. These results inform restoration strategies in arid‐region wetlands dominated by plants with slow litter decay where strategic litter removal may beneficially increase plant growth.