Spread of Exotic Cordgrasses and Hybrids (Spartina sp.) in the Tidal Marshes of San Francisco Bay, California, USA

Four species of exotic cordgrass (Spartina sp.) occur in the San Francisco estuary in addition to the California native Spartina foliosa. Our goal was to map the location and extent of all non-native Spartina in the estuary. Hybrids of S. alterniflora and S. foliosa are by far the most numerous exotic and are spreading rapidly. Radiating from sites of deliberate introduction, S. alterniflora and hybrids now cover ca. 190 ha, mainly in the South and Central Bay. Estimates of rate of aerial increase range from a constant value to an accelerating rate of increase. This could be due to the proliferation of hybrid clones capable of rapid expansion and having superior seed set and siring abilities. The total coverage of 195 ha by hybrids and other exotic cordgrass species is slightly less than 1% of the Bay's tidal mudflats and marshes. Spartina anglica has not spread beyond its original 1970s introduction site. Spartina densiflora has spread to cover over 5 ha at 3 sites in the Central Bay. Spartina patens has expanded from 2 plants in 1970 to 42 plants at one site in Suisun Bay. Spartina seed floats on the tide, giving it the potential to export this invasion throughout the San Francisco estuary, and to estuaries outside of the Golden Gate. We found isolated plants of S. alterniflora and S. densiflora in outer coast estuaries north of the Bay suggesting the likelihood for the San Francisco Bay populations to found others on the Pacific coast.     

Potential of Prokelisia spp. as Biological Control Agents of English Cordgrass, Spartina anglica

Spartina anglica, English cordgrass, has been isolated from North American herbivores since its 19th century amphidiploid hybrid origin in England. We found a population of this estuarine plant, introduced to Puget Sound, Washington, to be highly vulnerable to Prokelisia spp. planthoppers from California. More than 90% of plants died in greenhouse culture with high planthopper densities, while <1% died at the very low hopper densities of the control. English cordgrass invades the open intertidal mud of Pacific estuaries of North America, jeopardizing native flora and fauna of saltmarshes. This invasive plant alters the hydrology of marshes by accreting sediment. If intolerance to Prokelisia spp. in the field correlates with that in the greenhouse, biological control by this planthopper could contribute to management of cordgrass populations sensitive to this insect. Prokelisia spp. are stenophagous and pose little apparent threat to other plant genera. Neither the biological basis nor the inheritance of cordgrass vulnerability to the Prokelisia spp. are known. If genetic variability in vulnerability to this planthopper exists within the population of S. anglica, care would be needed to prevent increase of more resistant plant genotypes as a result of introduction of the insect.     

Variation in self-fertility and the reproductive advantage of self-fertility for an invading plant (Spartina alterniflora)

The factors responsible for the reproductive success or failure of individuals in small, founding populations have received little attention. Previous work on a small population of smooth cordgrass (Spartina alterniflora) invading San Francisco Bay, California found that most clones flower prolifically but set little or no seed, while a few clones have high rates of viable seed set, producing most of the seeds in the population. This study first identifies recruitment from seeds as the main source of new smooth cordgrass plants during invasion and then tests the influence of growing conditions and pollination treatment on viable seed set among clones established in San Francisco Bay. Field transplants indicated that a clone's seed set rate was not strongly dependent on its site of establishment. Low and high nutrient greenhouse treatments also had little effect on viable seed set rates within most clones. In contrast, pollination treatment (self-pollination or outcrossing) had a major effect on viable seed set rates. Most clones had high seed set rates after outcross-pollination, but clones varied widely in their selfing capacity. Zero or low viable self-seed set rates were most common; however, a few clones had high viable self-seed set rates, comparable to outcross seed set rates. A clone's selfing capacity was significantly correlated across years (r=0.89, P<0.001), and capacity to set viable self-pollinated seeds in the greenhouse was significantly correlated with the clone's rate of viable seed set in the field . In this growing population where cross-pollination is limited, only the clones with high selfing ability had high viable seed set rates in the field. Among primarily outcrossing plant invaders, variation in self-fertility among individuals may be a common phenomenon, with important implications for genetic differentiation, effective population size and patterns of spatial spread during an invasion.     

Assessing Invasion Impact: Survey Design Considerations and Implications for Management of An Invasive Marine Plant

We use a three-year study of sheltered low shore assemblages colonised by the non-indigenous Asian kelp Undaria pinnatifida to explore survey design issues for assessing the ecological impacts of invasive species. The weight of evidence overall suggested little impact from Undaria on low shore assemblages, with control–impact contrasts that could plausibly be interpreted as impacts probably reflecting natural causes. We demonstrate that the potential for reaching incorrect conclusions regarding the impacts of invasive species using control–impact designs is greater than when such designs are used to assess traditional forms of anthropogenic impact. We suggest that a before–after control–impact framework is essential, but recognise that such an approach has a number of limitations. In particular, there is no assurance that the before–after impact site will be invaded at all, or to the extent that provides worst-case impact information for coastal managers. We discuss possible ways of assessing invasive species impacts, but suggest that the uncertainty inherent in extrapolating impact information to other places and times means that the precautionary principle should be applied, and worst-case impacts assumed, until the level of scientific uncertainty is reduced. Such an approach should only be applied, however, after an evaluation of the feasibility, costs and benefits of managing a particular pest in relation to other priorities for invasive species.     

Simulating control strategies for a spatially structured weed invasion: <Emphasis Type="Italic">Spartina alterniflora</Emphasis> (Loisel) in Pacific Coast estuaries

Patterns in the spatial arrangement of invasive plant populations can provide opportunity for strategic placement of control efforts. Smooth cordgrass (Spartina alterniflora) is rapidly invading the intertidal mudflats of Pacific Coast estuaries. Its pattern of spread is distinctive. Seedlings establish in open mud and then grow vegetatively to form expanding circular patches, which dot the mudflats and eventually coalesce into a contiguous monospecific meadow. The invasion typically begins in the upper tide zone and then moves down the tidal gradient. A spatially explicit model was used to simulate the spread of S. alterniflora and compare various strategies for control in a situation where only a fraction of the total infestation could be controlled each year. A strategy of killing outlying patches first and then attacking the dense meadows (moving up the tidal gradient) led to eradication with up to 44% less time and effort than a strategy of attacking the dense meadows first and outlying patches second (moving down the tidal gradient). In the control of contiguous meadows located adjacent to the shoreline, the best strategy was to approach one end of the infestation, moving across the meadow to the other end. Suppression of seeds was not an effective control strategy. In general, effective control strategies were those that first eliminate the plant in areas where current or future vegetative growth is greatest. Application of these results in control programs for S. alterniflora and similar invasive species could greatly reduce the costs of control work and improve the likelihood of local or complete eradication.     

Effect of brassinolide on callus growth and regeneration in Spartina patens (Poaceae)

The effect of brassinolide (BL) on cultured calluses of Spartina patens (Ait.) Muhl. (Poaceae), a halophyte monocot was studied. BL at 0.03–0.04 mg l^–1 at fixed concentrations of IAA (0.2 mg l^–1) and BA (3.0 mg l^–1) added in MS medium increased the ratio for fresh weight (CIRFW) to dry weight (CIRDW) by 96–111% and 235–326%. Similarly, in callus regeneration capacity, BL at 0.03 mg l^–1 was most effective, increasing the shoot regeneration ratio (SRR) by 425%. BL at 0.04 mg l^–1 had not such an increasing effect as BL at 0.03 mg l^–1, which increased SRR by 79%. However, BL at 0.005 mg l^–1 promoted regenerated shoot growth most significantly, increasing the shoot height increasing ratio (SHIR) by 395% after a 40-day culture. BL at 0.05 mg l^–1 was least effective in the callus regeneration and regenerated shoot growth, decreasing SRR by 27% and SHIR by 52%. Present results suggest that BL at 0.03 mg l^–1 is suitable for the callus growth and shoot regeneration, while BL at 0.005 mg l^–1 effectively enhanced the regenerated shoot growth.     

Fungal biomass and decomposition in Spartina maritima leaves in the Mondego salt marsh (Portugal)

Spartina maritima (Curtis) Fernald is a dominant species in the Mondego salt marsh on the western coast of Portugal, and it plays a significant role in estuarine productivity. In this work, leaf litter production dynamics and fungal importance for leaf decomposition processes in Spartina maritima were studied. Leaf fall was highly seasonal, being significantly higher during dry months. It ranged from 42 g m^-2 in June to less than 6 g m^-2 during the winter. Fungal biomass, measured as ergosterol content, did not differ significantly between standing-decaying leaves and naturally detached leaves. Fungal biomass increased in wet months, with a maximum of 614 g g^-1 of ergosterol in January in standing-decaying leaves, and 1077 g g^-1 in December, in naturally detached leaves, decreasing greatly in summer. Seasonal pattern of fungal colonization was similar in leaves placed in litterbags on the marsh-sediment surface. However, ergosterol concentrations associated with standing-decaying and naturally detached leaves were always much higher than in litterbagged leaves, suggesting that fungal activity was more important before leaf fall. Dry mass of litterbagged leaves declined rapidly after 1 month (about 50%), mostly due to leaching of soluble organic compounds. After 13 months, Spartina leaves had lost 88% of their original dry weight. The decomposition rate constant (k) for Spartina maritima leaves was 0.151 month^-1.     

Blitzkrieg in a Marine Invasion: Caulerpa racemosa var. cylindracea (Bryopsidales, Chlorophyta) Reaches the Canary Islands (North-East Atlantic)

On the basis of morphological and genetic studies (rDNA ITS1, 5.8S, ITS2, and a 18S rDNA intron), we confirm here that Caulerpa racemosa var. cylindracea (Sonder) Verlaque, Huisman et Boudouresque, a southwestern Australian taxon recently introduced into the Mediterranean Sea also occurs in the Canary Islands. This is the first report of C. racemosa var. cylindracea in the Atlantic. It was observed for the first time in the Canary Archipelago in 1997–1998. The speed and regional scale of expansion (north Atlantic Ocean and the Mediterranean Sea) of this invasive species appear to be among the most dramatic ever recorded. The possible outcome of this introduction in the Atlantic is discussed.     

Oligochaeta in Spartina stems: the microdistribution of Enchytraeidae and Tubificidae in a salt marsh,Sapelo Island,USA

The distribution and abundance of Enchytraeidae and Tubificidae in and around Spartina alterniflora plants in a tidal salt marsh on Sapelo Island, Georgia, USA were studied using two different sampling techniques: wet funnel extraction and stem dissection. At least 80% of all worms inhabited leaf sheaths at the bases of S. alterniflora plants, and densities were low in sediment, root and surface debris samples. Oligochaete densities were dependent on the position within the marsh, the height on stems and the stage of sheath decay. Six predominant species were identified and included Marionina appendiculata, Marionina spartinae, Marionina waltersi, Marionina paludis, and Monopylephorus parvus. Individual species were distributed differently on stems and enchytraeids were more common than tubificids on standing-dead and further up S. alterniflora stems. Estimates of oligochaete densities in salt marsh habitats are increased dramatically when the numbers of worms on stems are considered. Possible advantages of the stem microhabitat are discussed in relation to the biology and ecology of oligochaetes.     

Modeling the Increase and Control of Caulerpa taxifolia, an Invasive Marine Macroalga

Population modeling based on species’ demography makes it possible to predict the pace of an invasion and evaluate the likelihood of success of different control strategies. We modeled the initial (density-independent) rate of increase of Caulerpa taxifolia (Vahl) C. Agardh (aquarium strain), a green alga that has markedly altered marine communities where it has invaded in the Mediterranean Sea. Parameter values for patch growth (from stolon extension) and reproduction (by asexual fragmentation and reattachment) were gleaned from published studies. Only the most conservative model, invoking field growth rates and low levels of fragment reattachment (2.5 m⁻² of existing patch each summer), closely matched observed increases (4–14 × annually). The most effective times for control (greatest reduction in rate of increase) were removal of established patches before summer and removal of fragments after summer. These times correspond to just before maximum growth and just after maximum reproduction, respectively. Only a combined strategy, incorporating 99% removal of all fragments and annual removal of 99% of established patches, was predicted to eliminate C. taxifolia entirely (λ < 1). This level of effort is only likely to be possible during the first few years of an invasion, arguing strongly for careful monitoring and rapid response to potential high-impact invaders.     

Invasion of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> Enhanced Ecosystem Carbon and Nitrogen Stocks in the Yangtze Estuary,China

Whether plant invasion increases ecosystem carbon (C) stocks is controversial largely due to the lack of knowledge about differences in ecophysiological properties between invasive and native species. We conducted a field experiment in which we measured ecophysiological properties to explore the response of the ecosystem C stocks to the invasion of Spartina alterniflora (Spartina) in wetlands dominated by native Scirpus mariqueter (Scirpus) and Phragmites australis (Phragmites) in the Yangtze Estuary, China. We measured growing season length, leaf area index (LAI), net photosynthetic rate (Pn), root biomass, net primary production (NPP), litter quality and litter decomposition, plant and soil C and nitrogen (N) stocks in ecosystems dominated by the three species. Our results showed that Spartina had a longer growing season, higher LAI, higher Pn, and greater root biomass than Scirpus and Phragmites. Net primary production (NPP) was 2.16 kg C m⁻² y⁻¹ in Spartina ecosystems, which was, on average, 1.44 and 0.47 kg C m⁻² y⁻¹ greater than that in Scirpus and Phragmites ecosystems, respectively. The litter decomposition rate, particularly the belowground decomposition rate, was lower for Spartina than Scirpus and Phragmites due to the lower litter quality of Spartina. The ecosystem C stock (20.94 kg m⁻²) for Spartina was greater than that for Scirpus (17.07 kg m⁻²), Phragmites (19.51 kg m⁻²) and the mudflats (15.12 kg m⁻²). Additionally, Spartina ecosystems had a significantly greater N stock (698.8 g m⁻²) than Scirpus (597.1 g m⁻²), Phragmites ecosystems (578.2 g m⁻²) and the mudflats (375.1 g m⁻²). Our results suggest that Spartina invasion altered ecophysiological processes, resulted in changes in NPP and litter decomposition, and ultimately led to enhanced ecosystem C and N stocks in the invaded ecosystems in comparison to the ecosystems with native species.     

Invasion Pattern of Herb Garlic Mustard (Alliaria petiolata) in High Quality Forests

The invasion of non-indigenous plant species poses a severe threat to native plant communities. Garlic mustard (Alliaria petiolata) is a naturalized European biennial herb that has spread rapidly through the eastern US and adjacent Canada. To determine garlic mustard rate of spread, eleven permanent plots (50×25 m) were located in seven high quality (relatively undisturbed) forests in the early stages of invasion. Garlic mustard presence was recorded within six 50×2 m permanent belt transects, and density and percent cover by age class were recorded in 36 permanent 1 m² quadrats, between 1989 and 1992, and again in 1997. Garlic mustard spread at an average rate of 5.4 m per year between 1989 and 1992, in all plots combined. Within individual plots rate of spread varied substantially, with location of the front increasing up to 36 m and decreasing as much as 18 m between years. While the front alternately advanced and retreated, over time garlic mustard consistently advanced through all forests. Rate of spread was influenced by establishment of satellite populations, and disturbance (wind-throw and flooding). The pattern of spread within plots was one of a ragged advancing front, supplemented by establishment of satellite populations 6–40 m distant from the front, which then coalesced with the main population. Garlic mustard presence between 1989 and 1997 increased significantly within all plots, and in each age class within each plot. The greatest increases occurred in plots where this plant was initially rarest. Garlic mustard cover and density varied nonsignificantly during the same time period. These results indicate that after garlic mustard invades a forest it becomes a permanent part of the community, annually increasing in presence but fluctuating in cover and density. Garlic mustard maintains a low profile under low disturbance conditions, but increases rapidly with periodic disturbance. This study monitored garlic mustard invasion in high quality relatively undisturbed forests, and may underestimate the rate of spread in low quality highly disturbed forests.     

Spatial and Temporal Dynamics of Exotic Earthworm Communities Along Invasion Fronts in a Temperate Hardwood Forest in South-Central New York (USA)

The invasion of North American forests by exotic earthworms is producing profound ecosystem changes, such as alterations in soil nutrient cycling, and redistribution and loss of soil organic matter. However, the present and future extent of these invasions is difficult to evaluate without a better understanding of the factors that control the distribution and abundance of earthworms in previously non-invaded habitats. In this study, the species composition and short-term dynamics of three exotic earthworm invasion fronts were studied at a northern hardwood forest in south-central New York State (USA). Belt transects were established at each of the three locations to sample from earthworm-invaded areas through transition zones and into invasion front areas. Lumbricus rubellus, L. terrestrisandOctolasion tyrtaeum were the most common species, but their distribution was not homogeneous along the transects. Whereas, L. rubellus was the only species with relatively high adult densities at transition zones and invasion fronts, L. terrestris and O. tyrtaeum occurred mostly in the heavily earthworm-invaded areas and were rare at the invasion fronts. The density of earthworms along the transects decreased by 60–87 from June 2001 to October 2002 and then recovered in 2003 to values similar to those of 2001. This decrease was apparently caused by reduced recruitment of immature earthworms, probably related to the severe drought periods that the study area experienced in 2001 and 2002. Our data suggest that climate and topography, through their effects on soil moisture patterns, can be critical factors controlling the distribution and spread of exotic earthworms in previously non-invaded habitats.     

Home and Away: Comparisons of Resource Utilization by a Marine Species in Native and Invaded Habitats

The exotic Asian shore crab, Hemigrapsus sanguineus, was recently introduced to the northeastern coast of North America and during the 1990's breeding populations were established throughout southern New England. In 1997–1998, ecological studies of several co-occurring brachyuran crabs were conducted and in native (Tanabe Bay, Japan) and invaded (Long Island Sound, USA) habitats of H. sanguineus. Standardized comparisons of H. sanguineus were made between the 2 habitats using data on crab sizes, utilization of space, and food habits. Results revealed that (1) the resource use of H. sanguineus was quite different from that of other resident species in its invaded habitat, and (2) there were no substantial changes in resource utilization by H. sanguineus after it became established in the invaded habitat (relative to native Tanabe Bay). Differing patterns of resource use by H. sanguineus and other crabs in the invaded habitat, the lack of restriction in resource use by H. sanguineus following its introduction, and the climatological and physical similarities between native and invaded regions likely contributed to the successful invasion of H. sanguineus into rocky intertidal habitats in southern New England.     

Evolutionary dynamics of Waxy and the origin of hexaploid Spartina species (Poaceae)

We investigated the evolutionary dynamics of duplicated copies of the granule-bound starch synthase I gene (GBSSI or Waxy) within polyploid Spartina species. Molecular cloning, sequencing, and phylogenetic analyses revealed incongruences between the expected species phylogeny and the inferred gene trees. Some genes within species were more divergent than expected from ploidy level alone, suggesting the existence of paralogous sets of Waxy loci in Spartina. Phylogenetic analyses indicate that this paralogy originated from a duplication that occurred prior to the divergence of Spartina from other Chloridoideae. Gene tree topologies revealed three divergent homoeologous sequences in the hexaploid S. alterniflora that are consistent with the proposal of an allopolyploid origin of the hexaploid clade. Waxy sequences differ in insertion–deletion events in introns, which may be used to diagnose gene copies. Both paralogous and homoeologous coding regions appear to evolving under selective constraints.     

First record of the sea anemone Diadumene lineata (Verrill 1871) associated to Spartina alterniflora roots and stems, in marshes at the Bahia Blanca estuary, Argentina

We report the occurrence of the orange-striped green anemone Diadumene lineata (Verrill 1871) (=Haliplanella lineata) in salt marshes at the Bahía Blanca Estuary for the first time in August 2005. We also found this species attached to roots and stems of Spartina alterniflora, an association that has never been registered before. After their determination, sampling was performed during a year to evaluate seasonal abundance of this sea anemone. Results showed that D. lineata was present through the whole year, indicating the existence of a stable population. All individuals sampled were found attached to roots or stems of S. alterniflora, with the higher abundances detected in summer. Further studies are necessary to precise the potential effects of this exotic sea anemone on salt marsh communities.     

Mobility of metals in salt marsh sediments colonised by <Emphasis Type="Italic">Spartina maritima</Emphasis> (Tagus estuary,Portugal)

Chemical associations of Zn, Pb, Cu, Co and Cd were determined using a sequential extraction procedure in sediments colonised by S. maritima in three salt marshes within the Tagus estuary: Rosário, Corroios and Pancas. Concentrations of these metals were also analysed in above- and belowground parts of Spartina maritima, as well as in sediments colonised by the plant. The highest metal concentrations in sediments were found in the marshes near the industrial and urban areas, whereas metal concentrations in plants were not significantly different among sites. This was thought to be a consequence of differences observed in metal bioavailability: Metals in Pancas, the least polluted location, were largely associated to easily accessible fractions for plant uptake, probably as a result of low organic matter content and high sandy fraction in sediments. S. maritima was able to induce the concentration of metals between its roots in the three salt marshes. The results obtained in this study indicate that S. maritima could be useful to induce phytostabilisation of metals in sediments, although the effectiveness to modify chemical associations is highly dependent on existing sediment parameters, and thus different results could be obtained depending on site characteristics. Guest editors: J. Davenport, G. Burnell, T. Cross, M. Emmerson, R. McAllen, R. Ramsay & E. Rogan Challenges to Marine Ecosystems     

New Potential for Classical Biological Control ofEucallipterus tiliae(Homoptera: Drepanosiphidae)

Biological control efforts in California directed against the linden aphid (Eucallipterus tiliae) have been predicated on the basis that the aphid is native to Europe. Although several European parasitoids have become established and attackE. tiliaein California, complete control has not been achieved; the aphid is a problem in Europe as well. New records ofE. tiliaefrom eastern Asia suggest that the aphid may be native there. This area has the greatest species diversity of both the host genus (Tilia) and the other linden-feeding drepanosiphid aphids. It is recommended that any further efforts for locating importable natural enemies be focused in this region.