<Emphasis Type="BoldItalic">r</Emphasis>-Selected Traits in an Invasive Population

I compared life-history traits and self-fertilization rates in greenhouse culture of native Spartina alterniflora with an invasive population from a low-density Pacific estuary to see whether plants in the novel r-selective regime exhibit early reproduction, greater self-compatibility and high reproductive effort putting individuals at greater risk of death. Plants were grown from seed collected from the Atlantic and Gulf coasts of North America and a population introduced approximately 100 years ago to Willapa Bay, Washington, USA that has expanded to cover over ca. 6000 hectares. Pacific marshes have almost no native emergent vascular plants in the intertidal habitat, offering a virtually empty niche for invasive S. alterniflora. The low-density plants at the leading edge of this rapidly expanding population suffer a severe Allee effect of greatly reduced fecundity, caused by pollen limitation, compared to high-density areas. I found 99% of the invasive plants initiated reproduction in the first year of this study, while only 52% of the native range plants did so in the first year, followed by 34% in the second and 14% of native plants had not initiated reproduction after three years. The invasive plants had more than twice the reproductive effort of the native plants. Plants with the highest reproductive effort in the first year of growth died regardless of coastal site of origin, although nearly all of those that died over two years were invasive Pacific plants (27%). The invaders set two times the seed as the natives under forced selfing, suggesting greater self-compatibility or reproductive effort. These results suggest either a founding and bottleneck event and/or the invasive population has evolved from the long-lived, predominantly self-incompatible, K-selected state of the probable invasive propagules originating in dense, competitive native marshes.Co-ordinating editor: I. Olivieri     

One hundred years of Spartina altemiflora (Poaceae) in Willapa Bay, Washington: random amplified polymorphic DNA analysis of an invasive population

Spartina altemiflora (smooth cordgrass) has invaded Willapa Bay, Washington, covering bare mud flats and seagrass beds. At current expansion rates it threatens to occupy most of the intertidal habitat in the bay within 40 years. Although a major mitigation effort is underway, nothing is known about the genetic structure of the Willapa Bay S. altemiflora population or the underlying causes for the dramatic increase in seed set that has occurred in the last 15 years. Random amplified polymorphic DNA (RAPD) analysis was assessed as a method of DNA fingerprinting to analyse the genetic history and current structure of the Willapa population. The results suggest that all S. alterniflora clones in Willapa Bay are descended from a single genet. Given the limited genetic variability expected in a clonally founded population, we discuss the utility of RAPDs as genetic markers to explore the large morphological and reproductive differences reported among Willapa clones. Cluster analysis of RAPD fragments was used to show relatedness among S. alteniflora genets and may be helpful in locating the most prolific clones for mitigation efforts. Overall, RAPDs show great potential in genetic analysis of S. altemiflora, particularly in invasive populations where isozyme variability is low or nonexistent.     

Genetic and historical evidence disagree on likely sources of the Atlantic amethyst gem clam Gemma gemma (Totten, 1834) in California

Aim Historical information about source populations of invasive species is often limited; therefore, genetic analyses are used. We compared inference about source populations from historical and genetic data for the oyster‐associated clam, Gemma gemma that invaded California from the USA Atlantic coast. Location Mid‐Atlantic (North Carolina, Maryland), Northeastern (New Jersey, New York, Massachusetts) and the California coasts (Elkhorn Slough, San Francisco Bay, Bolinas Lagoon, Tomales Bay, Bodega Harbor). Methods The documented history of transplantation of Eastern oysters to California was reviewed. Cytochrome c oxidase subunit I (COI) sequences from recent and archived clams were examined in a haplotype network. We used AMOVA to detect geographic genetic structure and a permutation test for significant reductions in diversity. Results Chesapeake Bay oysters were transplanted to New York prior to shipment to San Francisco Bay and from there to peripheral bays. Gemma in the Northeastern and Mid‐Atlantic regions were genetically differentiated. In California, populations in Bodega Harbor and Tomales Bay were genetically similar to those in the Mid‐Atlantic area while clams in San Francisco Bay, Elkhorn Slough and Bolinas Lagoon resembled populations in the Northeastern region. In California, genetic variation was not highest in San Francisco Bay despite greater magnitude of oyster plantings. Haplotypes varied over time in native and introduced populations. Main Conclusions Historical records and inferences from genetics agree that both Northeastern and Mid‐Atlantic regions were sources for Gemma in California. Only complex genetic hypotheses reconcile the strong segregation of haplotypes in California to the historical evidence of mixing in their proximate source (New York). These hypotheses include sorting of mixtures of haplotypes or selection in non‐native areas. Haplotype turnover in San Francisco and Massachusetts samples over time suggests that the sorting hypothesis is plausible. We suggest, however, that Gemma was introduced independently and recently to Tomales Bay and Bodega Harbor.     

Phylogeography and cryptic introduction of the ragworm Hediste diversicolor (Annelida,Nereididae) in the Northwest Atlantic

Many benthic marine invertebrates show striking range disjunctions across broad spatial scales. Without direct evidence for endemism or introduction, these species remain cryptogenic. The common ragworm Hediste diversicolor plays a pivotal role in sedimentary littoral ecosystems of the North Atlantic as an abundant prey item and ecosystem engineer, but exhibits a restricted dispersal capacity that may limit connectivity at both evolutionary and ecological time scales. In Europe, H. diversicolor is subdivided into cryptic taxa and genetic lineages whose distributions have been modified by recent invasions. Its origin in the northwest Atlantic has not been adequately addressed. To trace the age and origin of North American ragworm populations, we analyzed mtDNA sequence data (COI) from the Gulf of Maine and Bay of Fundy (n=73 individuals) and compared our findings with published data from the northeast Atlantic. Our results together with previous data indicate that two species of the H. diversicolor complex have independently colonized the northwest Atlantic at least three different times, resulting in two distinct conspecific assemblages in the Bay of Fundy and Gulf of Maine (respectively) that are different from the species found in the Gulf of St. Lawrence. North American populations had significantly lower genetic diversity compared with populations in the northeast Atlantic, and based on patterns of shared identity, populations in the Bay of Fundy originated from the Baltic Sea and North Sea. Populations from the Gulf of Maine were phylogenetically distinct and most likely originated from unsampled European populations. Analyses of the North American populations revealed patterns of post‐colonization gene flow among populations within the Gulf of Maine and Bay of Fundy. However, we failed to detect shared haplotypes between the two regions, and this pattern of complete isolation corroborates a strong phylogeographic break observed in other species.     

Differential population response of allocation,phenology, and tissue chemistry in <Emphasis Type="Italic">Spartina alterniflora</Emphasis>

Phenotypic variation within species is widespread among salt marsh plants. For Spartina alterniflora, the dominant species of low intertidal wetlands across the Altantic and Gulf coasts of the US, distinct phenological and morphological differences among populations from different latitudes have been found. To determine whether S. alterniflora plants from lower latitudes and those regenerated from Delaware tissue cultures would maintain differences from that of native plants, we conducted a field study in a natural salt marsh in Delaware, US. After two growing seasons, plant height, stem density, above- and belowground biomass, elemental composition, and nutrient resorption were measured. Natural variation in porewater salinity influenced physiological traits of Na⁺/K⁺ ratio regulation and nitrogen resorption efficiency similarly across populations. While plant height exhibited plasticity where populations tended to converge to a similar height, several other traits remained distinct. Delaware plants had a greater rate of rhizome growth than Georgia and Louisiana plants, which correlated with a greater magnitude of fall senescence. If traits such as seasonal translocation are plastic and can change with the length of the growing season, climate warming may alter belowground biomass production of S. alterniflora in wetlands of the mid-Atlantic.     

Biomass and aboveground production of four angiosperms in Cantabrian (N. Spain) salt marshes

New data of aboveground biomass and production of four angiosperms over a 12 month period for the Cantabrian Sea salt marshes (Bay of Biscay, N. Spain) are presented. Based on harvest methods, maximum aboveground total biomass values for Spartina maritima (Curtis) Fernald, Spartina alterniflora Loisel, Salicornia ramosissima J. Woods and Halimione portulacoides (L.) Aellen were 628, 1109, 480 and 1267 gm^-2, respectively. We conclude that although a slight latitudinal gradient in biomass is revealed in the data compiled with reference to some of the species studied, more work is neccesary in order to assess the potential productivity of these ecosystems on the coasts of Europe and/or to make comparisons with salt marshes of the American coasts. Annual net aerial primary production estimates using Smalley's method were: 296, 1160, 486 and 952 gm^-2yr^-1, for Spartina maritima, Spartina alterniflora, Salicornia ramosissima and Halimione portulacoides, respectively. These results together with turnover rate estimates point to the lack of vigour of the native S. maritima, while the exotic S. alterniflora, which seems to be spreading along the Cantabrian estuaries, behaves like a veritable pionner throughout the low marshes in this region.     

Variable reproductive output among clones of Spartina alterniflora (Poaceae) invading San Francisco Bay,California: the influence of herbivory,pollination, and establishment site

Spartina alterniflora has recently been introduced to San Francisco Bay, California, and is rapidly invading open mud flats, growing in circular patches that we found to be individual genetic clones. We collected spikelet samples from more than 200 clones and observed germination rates ranging from 0% to 59%, indicating substantial variation in reproductive output among clones. Several experiments were performed to explore the cause of variation. Pollination manipulations showed that S. alterniflora is outcrossing, but pollen supplements did not increase spikelet germination rates. Exclusion of the only insect herbivore (a phloem feeder, Prokelisia marginata) from developing inflorescences increased the proportion of spikelets containing seed, but failed to increase germinations per spikelet. Spikelets from Willapa Bay, Washington, grown free of insect herbivores, had germination rates similar to San Francisco Bay. These results suggest herbivory is not limiting reproductive output of S. alterniflora. Spikelet viability was not related to clone size; however, clones located lower in the intertidal or far up a drainage slough averaged fewer germinations per spikelet, suggesting clones in areas with lower genet density may have lower spikelet viabilities. Spikelet samples from different sections of clones growing across wide environmental ranges had similar rates of germination, suggesting some genetic influence on spikelet viability. Differential reproductive output among clones and the novel selective environment of San Francisco Bay are expected to cause gene frequency changes in this rapidly expanding population.     

Genetic admixture accelerates invasion via provisioning rapid adaptive evolution

Genetic admixture, the intraspecific hybridization among divergent introduced sources, can immediately facilitate colonization via hybrid vigor and profoundly enhance invasion via contributing novel genetic variation to adaption. As hybrid vigor is short‐lived, provisioning adaptation is anticipated to be the dominant and long‐term profit of genetic admixture, but the evidence for this is rare. We employed the 30 years' geographic‐scale invasion of the salt marsh grass, Spartina alterniflora, as an evolutionary experiment and evaluated the consequences of genetic admixture by combining the reciprocal transplant experiment with quantitative and population genetic surveys. Consistent with the documentation, we found that the invasive populations in China had multiple origins from the southern Atlantic coast and the Gulf of Mexico in the US. Interbreeding among these multiple sources generated a “hybrid swarm” that spread throughout the coast of China. In the northern and mid‐latitude China, natural selection greatly enhanced fecundity, plant height and shoot regeneration compared to the native populations. Furthermore, genetic admixture appeared to have broken the negative correlation between plant height and shoot regeneration, which was genetically‐based in the native range, and have facilitated the evolution of super competitive genotypes in the invasive range. In contrast to the evolved northern and mid‐latitude populations, the southern invasive populations showed slight increase of plant height and shoot regeneration compared to the native populations, possibly reflecting the heterotic effect of the intraspecific hybridization. Therefore, our study suggests a critical role of genetic admixture in accelerating the geographic invasion via provisioning rapid adaptive evolution.     

Multiple introductions and invasion pathways for the invasive ctenophore Mnemiopsis leidyi in Eurasia

The introduction and spread of non-indigenous species (NIS) in marine ecosystems accelerated during the twentieth century owing to human activities, notably international shipping. Genetic analysis has proven useful in understanding the invasion history and dynamics of colonizing NIS and identifying their source population(s). Here we investigated sequence variation in the nuclear ribosomal Internal Transcribed Spacer region of the ctenophore Mnemiopsis leidyi, a species considered one of the most invasive globally. We surveyed four populations from the native distribution range along the Atlantic coasts of the United States and South America, as well as six populations in the introduced range from the Black, Azov, Caspian and Baltic seas. Allelic and nucleotide diversity of introduced populations were comparable to those of native populations from which they were likely drawn. Introduced populations typically exhibited lower genetic differentiation (F _ST = ?0.014?C0.421) than native populations (F _ST = 0.324?C0.688). Population genetic analyses supported the invasion of Eurasia from at least two different pathways, the first from the Gulf of Mexico (e.g., Tampa Bay) to the Black Sea and thence to the Caspian Sea, the second from the northern part of the native distribution range (e.g., Narragansett Bay) to the Baltic Sea. The relatively high genetic diversity observed in introduced populations is consistent with large inocula and/or multiple invasions, both of which are possible given ballast water transport and the extensive native distribution of the ctenophore in the Atlantic Ocean.     

Foreword

Allozyme variation was characterised by starch gel electrophoresis at 23 enzyme coding loci and one regulatory locus in Atlantic salmon from 53 rivers in Eastern Canada, encompassing the majority of the species' North American range. Variation among rivers was highly heterogeneous and eight of the 15 polymorphisms showed regionally restricted distributions. Nearest neighbour joining (NJ) analysis and multi‐dimensional scaling suggest six distinct regional groups; Labrador/Ungava, Gulf of Saint Lawrence, Newfoundland (excluding Gulf rivers), the Atlantic shore/Southern Uplands of Nova Scotia, the inner Bay of Fundy, and the outer Bay of Fundy. Approximately 25% of observed genetic variation was distributed among these regions with a weak though significant overall correlation of genetic and geographic distance (Mantel Test, r = 0·255, P = 0·005). Collectively, the rivers showed consistent divergence from European populations with strong bootstrap support for the two clusters across loci in the NJ analysis. Mean heterozygosity was 0·061 for both continental groups, but the European population showed more than twice the variation among populations. F_ST values were 0·076 and 0·176 for North America and Europe, respectively, with an overall F_ST of 0·330.     

Genetically monomorphic invasive populations of the rapa whelk,Rapana venosa

Rapana venosa is a predatory marine gastropod native to the coastal waters of China, Korea, and Japan. Since the 1940s, R. venosa has been transported around the globe and introduced populations now exist in the Black Sea, the Mediterranean Sea, the Adriatic and Aegean seas, off the coasts of France and the Netherlands, in Chesapeake Bay, Virginia, USA, and in the Rio de la Plata between Uruguay and Argentina. This study surveyed variation in two mitochondrial gene regions to investigate the invasion pathways of R. venosa, identify likely sources for introduced populations, and evaluate current hypotheses of potential transportation vectors. Sequence data were obtained for the mitochondrial cytochrome c oxidase I and NADH dehydrogenase subunit 2 gene regions of 178 individuals from eight native locations and 106 individuals from 12 introduced locations. Collections from within the native range displayed very high levels of genetic variation while collections from all introduced populations showed a complete lack of genetic diversity; a single haplotype was common to all introduced individuals. This finding is consistent with the hypothesis that R. venosa was initially introduced into the Black Sea, and this Black Sea population then served as a source for the other secondary invasions by various introduction vectors including ballast water transport. Although non‐native R. venosa populations currently appear to be thriving in their new environments, the lack of genetic variability raises questions regarding the evolutionary persistence of these populations.     

Population structure of the dusky pipefish (Syngnathus floridae) from the Atlantic and Gulf of Mexico,as revealed by mitochondrial DNA and microsatellite analyses

Aim To elucidate the historical phylogeography of the dusky pipefish (Syngnathus floridae) in the North American Atlantic and Gulf of Mexico ocean basins. Location Southern Atlantic Ocean and northern Gulf of Mexico within the continental United States. Methods A 394‐bp fragment of the mitochondrial cytochrome b gene and a 235‐bp fragment of the mitochondrial control region were analysed from individuals from 10 locations. Phylogenetic reconstruction, haplotype network, mismatch distributions and analysis of molecular variance were used to infer population structure between ocean basins and time from population expansion within ocean basins. Six microsatellite loci were also analysed to estimate population structure and gene flow among five populations using genetic distance methods (F_ST, Nei’s genetic distance), isolation by distance (Mantel’s test), coalescent‐based estimates of genetic diversity and migration patterns, Bayesian cluster analysis and bottleneck simulations. Results Mitochondrial analyses revealed significant structuring between ocean basins in both cytochrome b (Φ_ST = 0.361, P < 0.0001; Φ_CT = 0.312, P < 0.02) and control region (Φ_ST = 0.166, P < 0.0001; Φ_CT = 0.128, P < 0.03) sequences. However, phylogenetic reconstructions failed to show reciprocal monophyly in populations between ocean basins. Microsatellite analyses revealed significant population substructuring between all locations sampled except for the two locations that were in closest proximity to each other (global F_ST value = 0.026). Bayesian analysis of microsatellite data also revealed significant population structuring between ocean basins. Coalescent‐based analyses of microsatellite data revealed low migration rates among all sites. Mismatch distribution analysis of mitochondrial loci supports a sudden population expansion in both ocean basins in the late Pleistocene, with the expansion of Atlantic populations occurring more recently. Main conclusions Present‐day populations of S. floridae do not bear the mitochondrial DNA signature of the strong phylogenetic discontinuity between the Atlantic and Gulf coasts of North America commonly observed in other species. Rather, our results suggest that Atlantic and Gulf of Mexico populations of S. floridae are closely related but nevertheless exhibit local and regional population structure. We conclude that the present‐day phylogeographic pattern is the result of a recent population expansion into the Atlantic in the late Pleistocene, and that life‐history traits and ecology may play a pivotal role in shaping the realized geographical distribution pattern of this species.     

Characterization of microsatellite loci in Spartina species (Poaceae)

The cordgrasses in the genus Spartina have become model organisms for studying biological invasions from both ecological and genetic perspectives. Here we characterize 11 disomic loci in Spartina alterniflora that show promise for population studies and for studying hybridization events between S. alterniflora and S. foliosa. Comparisons among invasive and native S. alterniflora populations showed that levels of allelic variation are lower in invasive populations. In addition, nearly all loci that amplified in S. foliosa populations and in a swarm of S. alterniflora×foliosa hybrids were polymorphic. We also found that several loci amplified successfully in other Spartina species.     

Assessing the geographic scale of genetic population management with microsatellites and introns in the clam Ruditapes decussatus

The clam Ruditapes decussatus is commercially important in southwestern Europe, suffering from population decline and hybridization with exotic Manila clam (R. philippinarum). Previous studies with intronic markers showed a genetic subdivision of the species in three races (Atlantic, West Mediterranean, and Adriatic‐Aegean). However, detailed population genetic studies to help management of the main production areas in the southwest of Europe are missing. We have analyzed eight Atlantic and two Mediterranean populations from the Spanish coasts using 14 microsatellites and six intronic markers. Microsatellites confirmed the Atlantic and West Mediterranean races detected with introns and showed that genetic variability was higher in Mediterranean than in Atlantic populations. Both marker types showed that genetic differentiation of Atlantic populations was low and indicated that populations could be managed at the regional level in the case of Cantabrian and Gulf of Cadiz areas, but not in the case of Rias Baixas and the Mediterranean. This study shows the interest of including different types of markers in studies of genetic population structure of marine organisms.     

Phenotypic plasticity of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> and <Emphasis Type="Italic">Phragmites australis</Emphasis> in response to nitrogen addition and intraspecific competition

Phenotypic plasticity of the two salt marsh grasses Spartina alterniflora and Phragmites australis in salt marshes is crucial to their invasive ability, but the importance of phenotypic plasticity, nitrogen levels, and intraspecific competition to the success of the two species is unclear at present. Spartina alterniflora Loisel. is an extensively invasive species that has increased dramatically in distribution and abundance on the Chinese and European coasts, and has had considerable ecological impacts in the regions where it has established. Meanwhile, Phragmites australis Cav., a native salt marsh species on the east coast of China, has replaced the native S. alterniflora in many marshes along the Atlantic Coast of the US. This study determined the effects of nitrogen availability and culm density on the morphology, growth, and biomass allocation traits of Spartina alterniflora and Phragmites australis. A large number of morphological, growth, and biomass parameters were measured, and various derived values (culm: root ratio, specific leaf area, etc.) were calculated, along with an index of phenotypic plasticity. Nitrogen addition significantly affected growth performance and biomass allocation traits of Spartina alterniflora, and culm density significantly affected morphological characteristics in a negative way, especially for Spartina alterniflora. However, there were no significant interactions between nitrogen levels and culm density on the morphological parameters, growth performances parameters, and biomass allocation parameters of the two species. Spartina alterniflora appears to respond more strongly to nitrogen than to culm density and this pattern of phenotypic plasticity appears to offer an expedition for successful invasion and displacement of Phramites australias in China. The implication of this study is that, in response to the environmental changes that are increasing nitrogen levels, the range of Spartina alterniflora is expected to continue to expand on the east coast of China.     

Exotic Spartina alterniflora provides compatible habitats for native estuarine crab Sesarma dehaani in the Yangtze River estuary

Although the impact of plant invasions on benthic communities, especially burrowing crabs, has received increasing attention, the results from past studies are mixed. The exotic plant Spartina alterniflora has become the most abundant species in the salt marshes of the Yangtze River estuary since it was first found just over a decade ago, but its effects on crabs in the salt marshes is largely unknown. To examine whether the invasions of this exotic plant affected native crabs, we compared the biomass and abundance of the dominant burrowing crab Sesarma dehaani in an exotic Spartina marsh, native Phragmites australis marsh and mudflats of the Yangtze River estuary, China. To explain the differences of S. dehaani populations between different habitats, feeding preference of S. dehaani for Spartina and Phragmites was investigated. Results showed crab abundance and biomass in the Spartina marsh were significantly greater than those in the Phragmites marsh and mudflats. Soil water content and plant community characteristics in the Spartina marsh also significantly differed in the Phragmites marsh and mudflats. Moreover, the feeding preference experiment showed that crabs consumed Spartina more than twice as much as Phragmites. In summary, this study showed that Spartina provided compatible habitats for native crab S. dehaani through offering suitable food source and moderate environmental conditions.     

Hybridization between <Emphasis Type="Italic">Schoenoplectus</Emphasis> sedges across Chesapeake Bay marshes

Hybridization is an evolutionary mechanism capable of enhancing adaptive potential, especially among species in fragmented or disturbed ecosystems like coastal marshes. In this study, we evaluated whether hybridization might influence adaptive responses in coastal marshes that are susceptible to the effects of global environmental change. To do so, we examined the extent and nature of hybridization between Schoenoplectus americanus and S. pungens, two ecologically dominant sedges in low-lying marshes across Chesapeake Bay (USA). Observed patterns of variation at genetically based morphological traits, cpDNA and nuclear microsatellite markers confirm that introgressive hybridization occurs between the two species. Comparisons of microsatellite and cpDNA profiles found that hybridization is reciprocal, although a disproportionate number of hybrids exhibit genomic asymmetries favoring S. americanus. AIC model selection consistently identified latitude as the strongest explanatory variable for the distribution of parental species, although discriminant analysis indicated that distributions also correspond to variation in environmental conditions. Discriminant analysis further indicated that ecological correlates of hybrid and S. americanus genotypes are similar, but not uniformly so. These findings indicate that the boundary between S. americanus and S. pungens is porous, and that hybridization could influence responses of one or both species to changing environmental regimes.     

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.     

Control and consequences of <Emphasis Type="Italic">Spartina</Emphasis> spp. invasions with focus upon San Francisco Bay

Maritime Spartina spp. are powerful ecosystem engineers that accrete sediment, define shorelines, create habitat, and generate prodigious primary productivity both where they are native and where they have been introduced. Invasive Spartina spp. can compete vigorously with native species, diminish biota, change hydrology, and confound human uses of estuaries. Herbicides have been effective in controlling several Spartina spp. invasions. One of the most recent successes is a 15-year campaign that has virtually eliminated S. alterniflora from the large, century-old invasion in Willapa Bay, WA, USA. Hybridization between native and introduced Spartina spp. has created new species and hybrid swarms. In San Francisco Bay, CA, USA (SF Bay) a complicated situation continues to play out from the purposeful introduction of S. alterniflora, which hybridized with native California cordgrass, S. foliosa. The hybrids spread rapidly and led to a long list of environmental problems, which led to an herbicide program that was successful in greatly diminishing the hybrid and saving the open mud habitat of migratory shorebirds. However, it was belatedly realized that the non-migratory, endangered Ridgeway’s rail uses the tall, dense hybrid Spartina as a surrogate for habitat that was lost during the twentieth century to urbanization and agricultural transformation of marshes around SF Bay. This realization has made difficult the simultaneous management of hybrid Spartina, wildlife conservation, and marsh restoration in San Francisco Bay. Restoration of native vegetation could satisfy the multiple goals of preserving open mud and conserving Ridgeway’s rail.