Does invasion of hybrid cordgrass change estuarine food webs?

Studies examining the impacts of introduced species on food webs often focus on the top-down effects of introduced predators. However, marine and estuarine systems have been invaded by plants that have the potential to alter carbon and nitrogen sources available to consumers. In San Francisco Bay, California, USA, hybridized cordgrass Spartina alterniflora × foliosa is adding C₄ carbon biomass to this system. We used natural abundances of stable isotopes of carbon and nitrogen to examine whether infaunal and epifaunal food webs reflected the large detrital input from hybrid Spartina. We compared stable isotope signatures among macrofaunal invertebrate consumers collected in hybrid Spartina, native S. foliosa, or unvegetated mudflats. We found no additional shift towards hybrid Spartina in hybrid areas. Structural changes brought about by an invasive ecosystem engineer, specifically increased biomass and detrital inputs, do not necessarily result in its increased incorporation into the food web.     

Influence of invasive <Emphasis Type="Italic">Spartina</Emphasis> growth stages on associated macrofaunal communities

In coastal wetlands, invasive plants often act as ecosystem engineers altering flow, light and sediments which, in turn, can affect benthic animal communities. However, the degree of influence of the engineer will vary significantly as it grows, matures and senesces, and surprisingly little is known about how the influence of an ecosystem engineer varies with ontogeny. We address this issue on the tidal flats of San Francisco Bay where hybrid Spartina (foliosa × alterniflora) invaded 30 years ago. The invasion has altered the physico-chemical properties of the sediment habitat, which we predicted should cause changes in macrofaunal community structure and function. Through mensurative and manipulative approaches we investigated the influence of different growth stages of hybrid Spartina on macrobenthos and the underlying mechanisms. Cross-elevation sampling transects were established covering 5 zones (or stages) of the invasion, running from the tidal flat (pre-invasion) to an unvegetated dieback zone. Additionally, we experimentally removed aboveground plant structure in the mature (inner) marsh to mimic the ’unvegetated areas’. Our results revealed four distinct faunal assemblages, which reflected Spartina-induced changes in the corresponding habitat properties along an elevation gradient: a pre-invaded tidal flat, a leading edge of immature invasion, a center of mature invasion, and a senescing dieback area. These stages of hybrid Spartina invasion were accompanied by a substantial reduction in macrofaunal species richness and an increase in dominance, as well as a strong shift in feeding modes, from surface microalgal feeders to subsurface detritus/Spartina feeders (mainly tubificid oligochaetes and capitellid polychaetes). Knowledge of the varying influence of plant invaders on the sediment ecosystem during different phases of invasion is critical for management of coastal wetlands.     

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.     

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.     

Tidal and seasonal effects on survival rates of the endangered California clapper rail: does invasive Spartina facilitate greater survival in a dynamic environment?

Invasive species frequently degrade habitats, disturb ecosystem processes, and can increase the likelihood of extinction of imperiled populations. However, novel or enhanced functions provided by invading species may reduce the impact of processes that limit populations. It is important to recognize how invasive species benefit endangered species to determine overall effects on sensitive ecosystems. For example, since the 1990s, hybrid Spartina (Spartina foliosa × alterniflora) has expanded throughout South San Francisco Bay, USA, supplanting native vegetation and invading mudflats. The endangered California clapper rail (Rallus longirostris obsoletus) uses the tall, dense hybrid Spartina for cover and nesting, but the effects of hybrid Spartina on clapper rail survival was unknown. We estimated survival rates of 108 radio-marked California clapper rails in South San Francisco Bay from January 2007 to March 2010, a period of extensive hybrid Spartina eradication, with Kaplan–Meier product limit estimators. Clapper rail survival patterns were consistent with hybrid Spartina providing increased refuge cover from predators during tidal extremes which flood native vegetation, particularly during the winter when the vegetation senesces. Model averaged annual survival rates within hybrid Spartina dominated marshes before eradication (? = 0.466) were greater than the same marshes posttreatment (? = 0.275) and a marsh dominated by native vegetation (? = 0.272). However, models with and without marsh treatment as explanatory factor for survival rates had nearly equivalent support in the observed data, lending ambiguity as to whether hybrid Spartina facilitated greater survival rates than native marshland. Conservation actions to aid in recovery of this endangered species should recognize the importance of available of high tide refugia, particularly in light of invasive species eradication programs and projections of future sea-level rise.     

The introduced ribbed mussel (<Emphasis Type="Italic">Geukensia demissa</Emphasis>) in Estero de Punta Banda,Mexico: interactions with the native cord grass, <Emphasis Type="Italic">Spartina foliosa</Emphasis>

Introduced populations of Guekensia demissa occur on the west coast of North America. They have been reported in San Francisco Bay, four southern California wetlands, and in Estero de Punta Banda (EPB), Baja California Norte, Mexico. We randomly sampled benthic invertebrates in four habitat types within EPB: marsh, channel, mudflat and pan. Geukensia demissa was the most abundant bivalve in the wetland at EPB. It was significantly associated with the native cordgrass, Spartina foliosa, and occurred at higher average densities in vegetated marsh sites (24/m²) and Spartina-dominated tidal channels (35/m²), compared to mudflat (0/m²), and pan (0/m²) sites. We estimated that the total biomass of this invader was over four times that of the next most abundant bivalve, Tagelus spp., in EPB. We examined G. demissa for parasites and found that only a few native parasites colonized this introduced host at very low prevalences and intensities. We performed bird surveys to determine the habitat overlap and potential impact of this mussel on the EPB population of light-footed clapper rails (Rallus longirostrus levipes), an endangered species in the United States. The high abundance of G. demissa in EPB, its presence in clapper rail habitat, and its known effects on salt marsh habitat in it’s native range, warrant further investigations of the impact of this invader in EPB and elsewhere.     

Ascomycete fungal communities associated with early decaying leaves of Spartina spp. from central California estuaries

Ascomycetous fungi play an important role in the early stages of decomposition of Spartina alterniflora, but their role in the decomposition of other Spartina species has not been investigated. Here we use fingerprint (terminal restriction fragment length polymorphism) and phylogenetic analyses of the 18S to 28S internal transcribed spacer region to compare the composition of the ascomycete fungal communities on early decay blades of Spartina species (Spartina alterniflora, Spartina densiflora, Spartina foliosa, and a hybrid (S. alterniflora × S. foliosa)) collected from three salt marshes in San Francisco Bay and one in Tomales Bay, California, USA. Phaeosphaeria spartinicola was found on all samples collected and was often dominant. Two other ascomycetes, Phaeosphaeria halima and Mycosphaerella sp. strain 2, were also common. These three species are the same ascomycetes previously identified as the dominant fungal decomposers on S. alterniflora on the east coast. Ascomycetes appeared to exhibit varying degrees of host specificity, demonstrated by grouping patterns on phylogenetic trees. Neither the exotic S. alterniflora nor the hybrid supported fungal flora different from that of the native S. foliosa. However, S. densiflora had a significantly different fungal community than the other species, and hosted at least two unique ascomycetes. Significant differences in the fungal decomposer communities were also detected within species (two clones of S. foliosa), but these were minor and may be due to morphological differences among the plants.     

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.     

Spatial patterns and species performances in experimental plant communities

The roles of co-occurring herbivores that modify habitat structure and ecosystem processes have seldom been examined in manipulative experiments or explored in early successional communities. In a created marsh in southern California (USA), we tested the individual and combined effects of two epibenthic invertebrates on nutrient and biomass pools, community structure, and physical habitat features. We manipulated snail (Cerithidea californica) and crab (Pachygrapsus crassipes) presence in field enclosures planted with pickleweed (Salicornia virginica) at elevations matching the plant’s lower extent in an adjacent natural marsh. In the 4-month experiment, C. californica altered habitat structure by reducing sediment surface heterogeneity and shear strength (a measure of sediment stability) markedly throughout the enclosures. Both invertebrates had strong negative effects on a group of correlated sediment physicochemical characteristics, including nitrogen and organic matter concentrations and soil moisture. In addition, both invertebrates greatly reduced benthic chlorophyll a, a proxy for biomass of microphytobenthos. Compared to controls, macroalgal cover was up to sixfold lower with crabs present, while snails increased cover at low elevations of enclosures. Unexpectedly, macroalgal cover was eliminated with both species present, perhaps through P. crassipes consumption of larger thalli and C. californica reduction in cover of recruits. Neither species influenced the S. virginica canopy (quantified with an index of branch length and number); however, at the lower elevation of enclosures, the two species together negatively impacted the plant canopy. The two invertebrates’ modifications to our experimental marshes led to distinct suites of biotic and physicochemical features depending on their presence or co-occurrence, with the latter producing several unexpected results. We propose that the roles and interactions of habitat-modifying fauna deserve further attention, particularly in the context of efforts to conserve and restore the processes found in natural systems.     

<Emphasis Type="Italic">Spartina maritima</Emphasis> influence on the dynamics of the phosphorus sedimentary cycle in a warm temperate estuary (Mondego estuary,Portugal)

During the last decades the Mondego estuary has been under severe ecological stress mainly caused by eutrophication. In this salt march system, Spartina maritima covers about 10.5 ha of the intertidal areas. The objective of the present study was to evaluate the effect of Spartina maritima marshes on the dynamics of phosphorus (P) binding in the surface sediment. We compare phosphate and oxygen fluxes, P-adsorption capacity, phosphate concentrations and total amount, and the extractable P forms in the upper 20 cm of sediment in vegetated sediment with adjacent mudflats without vegetation. Sediment pore-water profiles followed a clear trend, with lower P concentrations in more superficial layers, and increasing with depth. The vegetated mudflats presented lower concentrations of dissolved inorganic phosphorus than adjacent bare bottom mudflats, lower phosphate total amount, as well as higher P-adsorption capacity. Results from the extraction procedure show that the superficial layers are the most important for estuarine phosphorus dynamics, since maximum concentrations of labile P pools are present here. In contrast, higher proportions of refractory P pool are found in deeper layers. Spartina marsh sediments had less total P, less iron bound P, and less exchangeable P than adjacent bare bottom mudflats. Also the pool of loosely sorbed P is lower in the Spartina marsh. Phosphate regeneration from the sediment to the overlying water was only 11.8 kg ha⁻¹ year⁻¹ in vegetated sediment while 25.8 kg ha⁻¹ year⁻¹ in the bare mud flat. Plant uptake for growth combined with an enhanced P-adsorption capacity of the sediment, may explain these differences. Therefore, Spartina marshes are very important agents in the sedimentary P cycle worldwide, and can be considered a useful management tool in estuarine ecosystem recovery efforts.     

Food web analysis of southern California coastal wetlands using multiple stable isotopes

Carbon, nitrogen, and sulfur stable isotopes were used to characterize the food webs (i.e., sources of carbon and trophic status of consumers) in Tijuana Estuary and San Dieguito Lagoon. Producer groups were most clearly differentiated by carbon, then by sulfur, and least clearly by nitrogen isotope measurements. Consumer ¹⁵N isotopic enrichment suggested that there are four trophic levels in the Tijuana Estuary food web and three in San Dieguito Lagoon. A significant difference in multiple isotope ratio distributions of fishes between wetlands suggested that the food web of San Dieguito Lagoon is less complex than that of Tijuana Estuary. Associations among sources and consumers indicated that inputs from intertidal macroalgae, marsh microalgae, and Spartina foliosa provide the organic matter that supports invertebrates, fishes, and the light-footed clapper rail (Rallus longirostris levipes). These three producers occupy tidal channels, low salt marsh, and mid salt marsh habitats. The only consumer sampled that appears dependent upon primary productivity from high salt marsh habitat is the sora (Porzana carolina). Two- and three-source mixing models identified Spartina as the major organic matter source for fishes, and macroalgae for invertebrates and the light-footed clapper rail in Tijuana Estuary. In San Dieguito Lagoon, a system lacking Spartina, inputs of macroalgae and microalgae support fishes. Salicornia virginica, S. subterminalis, Monanthochloe littoralis, sewage- derived organic matter, and suspended particulate organic matter were deductively excluded as dominant, direct influences on the food web. The demonstration of a salt marsh–channel linkage in these systems affirms that these habitats should be managed as a single ecosystem and that the restoration of intertidal marshes for endangered birds and other biota is compatible with enhancement of coastal fish populations; heretofore, these have been considered to be competing objectives. Received: 24 April 1996 / Accepted: 24 October 1996     

Impacts of Spartina alterniflora invasion on the benthic communities of salt marshes in the Yangtze Estuary,China

As a species for ecological engineering, Spartina alterniflora was introduced to Chongming Dongtan in 1995, and over the last 10 years, this species has rapidly invaded large areas of the Chongming Dongtan nature reserve. In this study, use of a normalized biomass size-spectra (NBSS) approach was explored to evaluate the possible impacts of S. alterniflora invasion on the benthic communities along gradients of intertidal zones and the invasion history of S. alterniflora within the nature reserve. The results showed that the characteristics of macrobenthic communities and the variation in macrobenthic communities described by the first two CCA axes revealed clearly the gradients of elevation and invasion history of S. alterniflora. The differences in the macrobenthic assemblages between the Spartina alterniflara marshes and the native Phragmites australis marshes decreased with increasing of invasion history of S. alterniflara. The macrobenthic biomass showed a decreasing trend, while the meiobenthic biomass showed a reverse trend along the elevation gradient. The macrobenthic biomass of S. alterniflora marshes with longer invasion history was higher than that at recently invaded S. alterniflora marshes, while the meiobenthic biomass was lower. The slopes of NBSS for the sampling sites showed a trend of steeper slopes with decreasing of elevation and at the recently invaded S. alterniflora marshes than that at marshes with longer invasion history, while the differences between the native P. australis marshes and the S. alterniflora marshes with long invasion history tended to be diminished. The NBSS approach could thus be used more widely to detect possible impacts of S. alterniflara invasion on benthic assemblages. This study also indicated the potential for this approach to provide valuable insights into the ecosystem ecology of invasive species, which could be very important for wetland biodiversity conservation and resource management in the Yangtze River Estuary and other such impacted areas.     

Effects of a habitat-altering invader on nesting sparrows: An ecological trap?

Many invading species impact native species through predation, parasitism or competition, while others affect natives indirectly by restructuring their habitat. How invasive plants affect native animals, and to what extent native animals respond to changes in their habitat and the novel selection pressures that follow, is not well known. We investigated the impacts of a habitat-altering invader, the Atlantic cordgrass Spartina alterniflora, on the nesting success of Alameda song sparrows (Melospiza melodia pusillula), a California Species of Special Concern, in tidal marshes in three sites in San Francisco Bay. Date of laying was the most influential factor in determining daily survival rate of nests, but whether the nest was placed in exotic Spartina was the most important ecological variable. Nests placed in exotic Spartina had a success rate that was 30% lower than those placed in native vegetation. Nests in exotic Spartina were significantly more likely to fail due to tidal flooding than were nests placed in native vegetation, because the densest stands of exotic Spartina occurred at significantly lower elevations relative to the tides. Our results suggest that exotic Spartina may be an ecological trap for song sparrows in San Francisco Bay, attracting birds to nest sites that are often destroyed by tidal flooding.     

Restoration of Arthropod Assemblages in a Spartina Salt Marsh following Removal of the Invasive Plant Phragmites australis

Invasive plants are one of the most serious threats to native species assemblages and have been responsible for the degradation of natural habitats worldwide. As a result, removal of invasive species and reestablishment of natural vegetation have been attempted in order to restore biodiversity and ecosystem function. This study examined how native arthropod assemblages, an abundant and functionally important group of organisms in many ecosystems, are affected by the incursion of the invasive wetland plant Phragmites australis and if the restoration of the native vegetation in brackish Spartina alterniflora marshes results in the reestablishment of the arthropod community. The invasion of Phragmites into a coastal Spartina marsh in southern New Jersey seriously altered arthropod assemblages and trophic structure by changing the abundance of trophic groups (detritivores, herbivores, carnivores) and their taxonomic composition. Herbivore assemblages shifted from the dominance of external free‐living specialists (e.g., planthoppers) in Spartina to concealed feeders in Phragmites (stem‐feeding cecidomyiids). Moreover, free‐living arthropods in Phragmites became dominated by detritivores such as Collembola and chironomids. The dominant marsh spiders, web‐building linyphiids, were significantly reduced in Phragmites habitats, likely caused by differences in the physical environment of the invaded habitats (e.g., lower stem densities). Thus, trophic structure of arthropod assemblages in Phragmites, as seen in the large shifts in feeding guilds, was significantly different from that in Spartina. Removal of Phragmites with the herbicide glyphosate resulted in the rapid return of Spartina (≤5 yrs). Moreover, return of the dominant vegetation was accompanied by the recovery of most original habitat characteristics (e.g., live and dead plant biomass, water flow rate). The arthropod assemblage associated with Spartina also quickly returned to its preinvasion state and was not distinguishable from that in uninvaded Spartina reference sites. This study provides evidence that the reestablishment of native vegetation in areas previously altered by an invasive plant can result in the rapid recovery of the native arthropod assemblage associated with the restored habitat.     

Nitrogen sequestration capacity of two salt marshes from the Tagus estuary

The role of salt marshes as nitrogen sink is examined taking into consideration the seasonal variation of above and belowground biomass of Spartina martima and Halimione portulacoides in two marshes from Tagus estuary, Pancas and Corroios, and the degradation rates of belowground litter. Total nitrogen was determined in plant components, decomposing litter and sediment. Biomass was higher in Corroios, the saltier marsh, with 7190 g m⁻² y⁻¹ dw of S. maritima and 6593 g m⁻² y⁻¹ dw of H. portulacoides and the belowground component contributed to 96% and 90% of total biomass, respectively. In the other marsh, Pancas, belowground biomass contributed to 56% and 76% of total biomass for S. maritima and H. portulacoides, respectively. Litterbag experiment showed that between 25% and 50% of nitrogen is lost within the first month and remained relatively constant in the next four months. Slower decomposition is observed in sediments with higher nitrogen concentration (max. 0.7% N in the saltier marsh). Higher concentrations of N were found in the sediment upper layers. Considering the sediment-root system, most of the nitrogen is stored in the sediment compartment and only about 1–4% of the total N was found in the roots. Considering these results, Tagus salt marshes act as a sink for nitrogen.     

Spartina alterniflora invasions in the Yangtze River estuary,China: An overview of current status and ecosystem effects

The Yangtze River estuary is an important ecoregion. However, Spartina alterniflora, native to North America, was introduced to the estuary in the 1990s through both natural dispersal and humans and now it is a dominant species in the estuarine ecosystems, with its invasions leading to multiple consequences to the estuary. S. alterniflora had great competitive effects on native species, including Scirpus mariqueter and Phragmites australis, and could potentially exclude the natives locally. The presence of S. alterniflora had little influence on the total density of soil nematodes and macrobenthonic invertebrates, but significantly altered the structure of trophic functional groups of nematode and macrobenthonic invertebrate communities. The conversion of mudflats to Spartina meadows had significant effects on birds of Charadriidae and Scolopacidae, which might be attributable to the reduction of food resources and the physical alterations of habitats for shorebirds. S. alterniflora invasions increased the primary productivity of the invaded ecosystems, and altered carbon and nitrogen cycling processes. Our studies focused mainly on the effects of S. alterniflora invasions on the structure of native ecosystems; thus further studies are clearly needed to investigate how ecosystem functioning is affected by the modification of the structure of estuarine ecosystems by S. alterniflora invasions.     

Nitrogen addition does not reduce belowground competition in a salt marsh clonal plant community in Mississippi (USA)

Previous studies have suggested that belowground competition for nutrients influences plant zonation in salt marshes. In this study, I tested the hypothesis that competition for nitrogen structured a clonal plant community in a nitrogen-limited salt marsh in coastal Mississippi, USA. In contrast to most previous field studies that have investigated mechanisms of competition, I examined clonal growth responses of established genets of a nitrogen-demanding low-intertidal species (Spartina alterniflora) to nitrogen addition and the removal of a nitrogen-conserving high-intertidal species (Juncus roemerianus). Nitrogen addition stimulated clonal invasion of the Juncus zone by Spartina but did not reduce the significant competitive effects of Juncus on Spartina. Simulated Juncus shade did not reduce invasion of the Juncus zone by Spartina, indicating that belowground competition reduced clonal invasion. In the last year of the study, the border shifted unexpectedly towards the Spartina zone, resulting in competitive displacement of Spartina by Juncus. Nitrogen addition did not prevent or slow this displacement, further contradicting the nitrogen competition hypothesis. Although growth rates were much more strongly limited by nitrogen in Spartina than in Juncus, nitrogen addition did not cause the displacement of Juncus by Spartina after three growing seasons. I conclude that zonation of Spartina and Juncus is maintained by preemption of space and greater tolerance of low nitrogen supplies by Juncus in the high marsh. These results contrast sharply with findings of reduced belowground competition with nutrient addition in previous studies and highlight the important role of nutrient-mediated competition for space between clonal plants.     

The rapid evolution of self-fertility in Spartina hybrids (Spartina alterniflora × foliosa) invading San Francisco Bay, CA

Plant hybridization can lead to the evolution of invasiveness. We wished to determine whether hybrids between the largely self-sterile Atlantic Spartina alterniflora and California native S. foliosa had evolved self-fertility during their ca 30 year existence in San Francisco Bay, CA. In pollination experiments we found that neither of the parental species was self-fertile, nor were early generation hybrids. A large fraction of later generation hybrids were profusely self-fertile. Inbreeding depression was high in the parental species and early generation hybrids, but was much reduced in later generation hybrids—some even showed outbreeding depression. We found that populations of later generation hybrids and their seedling progeny were almost two-fold more homozygous than early generation hybrids, consistent with the evidence of increased selfing shown by our parentage analyses based upon 17 microsatellite markers. We posit that evolved self-fertility has contributed substantially to the rapid spread of hybrid Spartina in San Francisco Bay.