The austral cordgrass Spartina densiflora Brong.: its taxonomy, biogeography and natural history

Aim During the last 20 years, the austral cordgrass Spartina densiflora has been recorded aggressively invading estuarine environments in the USA, Spain and Morocco. Whereas this species is one of the three most widely distributed worldwide, it is among the least studied within the genus. The objective of this work is to integrate baseline information about the taxonomy, global distribution, centre of origin, and general ecology of S. densiflora in native and invaded marshes worldwide in order to help to strengthen management efforts currently directed at controlling or eradicating it from locations where it has been introduced. Location World‐wide. Methods I review, update and discuss relevant information about S. densiflora published in peer‐reviewed papers, including those in journals with limited international distribution. I also review theses and major technical reports containing critical up‐to‐date information. Results This work indicates that, although S. densiflora remains in need of thorough scientific attention, key information on its taxonomy, distribution and invasive biology has been overlooked because it was published in languages other than English, and/or in journals with restricted distribution. Main conclusions Spartina densiflora seems to have originated along the east coast of South America; today, however, many other regions worldwide serve as donors for this invasive species, including Chile, the USA, Spain and Morocco. Spartina densiflora is a bioengineer organism, tolerant of a broad spectrum of environmental conditions and able to re‐shape the structure of invaded communities not just in mudflats, but also on sandy, muddy, and rocky shores as well as on cobble beaches. Only by integrating local‐scale research conducted in different geographical regions will we be able to understand the between‐site variations of its biological cycle, which in turn will aid in the design of more effective conservation strategies.     

Distribution of saltmarsh plant communities associated with environmental factors along a latitudinal gradient on the south‐west Atlantic coast

Aim To produce an inventory of south‐west Atlantic saltmarshes (from latitude 31°48′ S to 43°20′ S) using remotely sensed images and field sampling; to quantify their total area; to describe the biogeographical variation of the main habitats characterized by dominant vascular plants, in relation to major environmental factors; to test the hypothesis of predominance of the reversal pattern in plant distribution (sedges and grasses dominate the lower, regularly inundated zones, while the upper zones are occupied by more halophytic species) previously described; and to compare these south‐west Atlantic saltmarshes with others world‐wide. Location South‐western Atlantic saltmarshes Methods Field samples of dominant emergent plant species positioned by the global positioning system (GPS) were obtained from most coastal saltmarshes (14) between southern Brazil and northern Patagonia, Argentina. Landsat satellite images were obtained and coastal saltmarsh habitats were quantified by supervised classification, utilizing points gathered in the field. Results Three main plant species dominated the low and middle intertidal saltmarsh, Spartina alterniflora Loesel., Spartina densiflora Brong. and Sarcocornia perennis (P. Mill.) A.J. Scott. The total area of the studied coastal saltmarshes was 2133 km², comprising 380 km² of Sp. alterniflora marsh, 366 km² of Sp. densiflora marsh, 746 km² of Sar. perennis marsh and 641 km² of brackish marsh (dominated by Juncus acutus L., Juncus kraussii Hochst., Scirpus maritimus L., Scirpus americanus Pers. and Phragmites australis (Cav.) Trin.). Cluster analysis showed three habitat types: saltmarshes dominated by (1) Sp. densiflora and brackish species,(2) Sp. alterniflora and Sar. perennis and (3) Sp.densiflora only. The analysis of abiotic variables showed significant differences between groups of habitats and coordinated gradients of the abiotic variables. The south‐west Atlantic coast showed decreasing mean annual rainfall (1200 to 196 mm) and increasing mean tidal amplitude (< 0.5 to > 2.5 m) from latitude 31° to 43°. Main conclusions South‐west Atlantic saltmarshes are globally important by virtue of their total extent. Remote sensing showed that the reversal pattern in plant distribution is not widespread. Indeed, south‐west Atlantic saltmarshes are better characterized by the presence of the halophytic genera Spartina and Sarcocornia. Our results support the interpretation that south‐west Atlantic saltmarshes constitute a class of temperate type (sensu Adam, 1990 ) with transitional characteristics between Australasian–South African saltmarshes and west Atlantic saltmarshes.     

Positive interactions of the smooth cordgrass Spartina alterniflora on the mud snail Heleobia australis, in South Western Atlantic salt marshes

The role of positive interactions is often crucial in communities with intense abiotic stress such as intertidal environments. Grasses acting as ecosystem engineers, for example, may ameliorate intertidal harsh physical conditions and modify the community structure. The mud snails Heleobia australis d'Orbigny frequently inhabit the SW Atlantic marshes, mainly associated to intertidal marsh plants (mainly the smooth cordgrass Spartina alterniflora Loisel) probably due to the plant indirect effects. The purpose of this work was to investigate the magnitude of these association and the processes that generate the pattern. Samples of the snail abundance in six SW Atlantic coastal marshes show that H. australis is associated to coastal areas of low energy and low or none freshwater input. This result is important because this species is being used as bioindicator of coastal estuarine systems during the Holocene. Thus the paleontological interpretation based on this species should be revised. Within the studied areas, snails are associated to intertidal marsh plants. However, stable isotope analysis shows that neither plant nor their epiphytes are their main food sources. Field experiments show that snails actively select areas with plants, although tethering experiments show that plants do not provide shelter from predators. However, plants do buffer physical stress factors such as temperature, which generate important mortality outside plants covered areas. These positive interactions have large effects on H. australis distributions in marsh communities; increasing the habitats available for colonization and affecting their local distribution.     

芦苇、互花米草的生长和繁殖对盐分胁迫的响应

对2种海滨植物——土著种芦苇(Phragmites australis)和外来种互花米草(Sparti-na alterniflora)在淡水、中盐度(15‰)和高盐度(30‰)环境下生长和有性繁殖特征及其年际动态进行了研究,以期探索海滨植物对盐分胁迫的适应机制。结果表明:2年中各盐度处理下互花米草地上部分生物量均高于芦苇,芦苇和互花米草地上部分生物量、株高和分蘖数均随着盐度的升高而显著下降。芦苇第1年没有开花;第2年,只有淡水和中盐度处理下的芦苇开花。各盐分梯度下,第1年互花米草的开花株数没有显著差异,但花序重量在高盐度条件下显著下降;第2年高盐胁迫仍然抑制了互花米草的繁殖。第1年各处理下的芦苇株高均低于互花米草,但第2年淡水和中盐环境中的芦苇株高高于互花米草。除了高盐处理下的芦苇,第2年这2种植物地上部分生物量、株高、开花株数、花序重量显著高于第1年。与芦苇相比,互花米草表现出较高的生长能力和耐盐能力;2种植物的生长和繁殖的能力随着定居时间的增加而提高,但高盐胁迫抑制了芦苇第2年的增长;芦苇第2年在淡水和中盐度条件下表现出株高上的优势。     

Screening for bacterial-fungal associations in a south-eastern US salt marsh using pre-established fungal monocultures

Both bacteria and fungi play critical roles in decomposition processes in many natural environments, yet only rarely have they been studied as an integrated community. We examined whether physical associations exist between individual bacterial and fungal species that co-occur on decaying smooth cordgrass, Spartina alterniflora, in a south-eastern US salt marsh. Fungal-pervaded decaying Spartina was used as "bait" for potential bacterial associates. The bundles (infiltrated with one of three dominant fungal members of the decomposer assemblage, or an autoclaved control) were placed in a salt marsh and collected biweekly for 6 weeks during the first experiment (late summer 2002), and weekly for 3 weeks during the second experiment (early summer 2003). Terminal-restriction fragment length polymorphism (T-RFLP) analysis of 16S rRNA genes was used to track colonization by bacterial taxa in association with the established fungal species. T-RFLP analysis of 18S-to-28S internal transcribed spacer (ITS) regions was used to monitor changes in fungal communities once bundles had been placed in the field. Results from both years were nearly identical, and showed that invasion by fungi other than the bait species was slow, resulting in a virtual fungal monoculture for several weeks into the experiments. Surprisingly, bacterial communities were unaffected by the identity of the fungal bait. Regardless of the fungal species, and even in the absence of prior fungal colonization, bacterial 16S rRNA profiles were remarkably similar. These results suggest that few species-specific associations, either positive or negative, exist between bacterial and fungal members of the Spartina decomposer community during initial colonization.     

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.     

Functional characterization of phosphoenolpyruvate carboxykinase-type C4 leaf anatomy: immuno-, cytochemical and ultrastructural analyses

BACKGROUND AND AIMS: Species having C4 photosynthesis belonging to the phosphoenolpyruvate carboxykinase (PEP-CK) subtype, which are found only in family Poaceae, have the most complex biochemistry among the three C4 subtypes. In this study, biochemical (western blots and immunolocalization of some key photosynthetic enzymes) and structural analyses were made on several species to further understand the PEP-CK system. This included PEP-CK-type C4 species Urochloa texana (subfamily Panicoideae), Spartina alterniflora and S. anglica (subfamily Chloridoideae), and an NADP-ME-type C4 species, Echinochloa frumentacea, which has substantial levels of PEP-CK. KEY RESULTS: Urochloa texana has typical Kranz anatomy with granal chloroplasts scattered around the cytoplasm in bundle sheath (BS) cells, while the Spartina spp. have BS forming long adaxial extensions above the vascular tissue and with chloroplasts in a strictly centrifugal position. Despite some structural and size differences, in all three PEP-CK species the chloroplasts in mesophyll and BS cells have a similar granal index (% appressed thylakoids). Immunolocalization studies show PEP-CK (which catalyses ATP-dependent decarboxylation) is located in the cytosol, and NAD-ME in the mitochondria, in BS cells, and in the BS extensions of Spartina. In the NADP-ME species E. frumentacea, PEP-CK is also located in the cytosol of BS cells, NAD-ME is very low, and the source of ATP to support PEP-CK is not established. CONCLUSIONS: Representative PEP-CK species from two subfamilies of polyphyletic origin have very similar biochemistry, compartmentation and chloroplast grana structure. Based on the results with PEP-CK species, schemes are presented with mesophyll and BS chloroplasts providing equivalent reductive power which show bioenergetics of carbon assimilation involving C4 cycles (PEP-CK and NAD-ME, the latter functioning to generate ATP to support the PEP-CK reaction), and the consequences of any photorespiration.     

Genotypic diversity enhances invasive ability of Spartina alterniflora

Although genetic diversity is very important for alien species, which have to cope with new environments, little is known about the role that genetic diversity plays in their invasive success. In this study, we set up a manipulation experiment including three levels of genotypic diversity to test whether genotypic diversity can enhance the invasive ability of alien species, in our case the invasive Spartina alterniflora in China, and to infer the underlying mechanisms. There was no significant relationship between genotypic diversity and parameters of performance in the first year; however, from the summer of the second year onwards, genotypic diversity enhanced four of the six parameters of performance. After two growing seasons, there were significant positive relationships between genotypic diversity and maximum spread distance, patch size, shoot number per patch, and aboveground biomass. Moreover, abundance of the native dominant species Scirpus mariqueter was marginally significantly decreased with genotypic diversity of S. alterniflora, suggesting that enhanced invasive ability of S. alterniflora may have depressed the growth of the native species. There was no significant difference in most measures of performance among six genotypes, but we observed a transgressive over performance in four measures in multiple‐genotype patches. At the end of the experiment, there were significant nonadditive effects of genotypic diversity according to Monte Carlo permutations, in six‐genotype, but not three‐genotype plots. Our results indicated that both additive and nonadditive effects played roles in the positive relationship between genetic diversity and invasion success, and nonadditive effects were stronger as duration increased.     

镉在互花米草中积累、转运及亚细胞的分布

研究了在不同Cd浓度(0、5、100、200μg·g-1)处理下,互花米草花序、叶、茎、根茎、须根中Cd含量、积累量、转运特征,及Cd在互花米草体内的亚细胞分布。结果表明,Cd在互花米草不同器官中的积累能力存在较大差异。茎、根茎、须根中Cd含量及积累量随处理浓度的增加而升高,其中须根中Cd含量及积累量均高于其他器官。Cd处理浓度为100gμ·g-1时,花序和叶中Cd含量达到最大值,分别为8.65和7.82μg·g-1。在Cd处理浓度为200μg·g-1时,须根中Cd含量可高达390.00μg·g-1,积累量达3200μg·株-1。Cd在互花米草体内转运能力极低,绝大部分Cd积累在地下部位。Cd在互花米草亚细胞中的分布规律为细胞壁>胞液>细胞器。随着Cd处理浓度的增加,Cd在细胞壁中的分配比例增大,胞液中Cd分布比例则相应减小,细胞壁和胞液相互协调,增强互花米草对重金属Cd的耐性。     

Interactions between a hunting spider and a web-builder: consequences of intraguild predation and cannibalism for prey suppression

Abstract. 1. Antagonistic interactions among invertebrate predators such as intraguild predation and cannibalism have the potential to dampen top‐down impacts on shared prey at lower trophic levels. Two abundant spider predators, the large wolf spider Pardosa littoralis and the small sheet‐web builder Grammonota trivitatta co‐occur on the salt marshes of eastern North America where they both attack planthoppers (Prokelisia spp.), the dominant herbivores on the marsh. Experiments both in the laboratory and field were used to assess the incidence of intraguild predation and cannibalism in these spiders and elucidate how such antagonistic interactions influence planthopper suppression. 2. Functional response experiments showed that with an increase in planthopper prey density, Grammonota captured more prey but not a higher proportion of that offered. Pardosa exhibited the same response when Grammonota were offered as intraguild prey. Both functional responses were type I over the range of prey densities offered. 3. Grammonota is moderately cannibalistic, and the presence of planthopper prey reduced the incidence of cannibalism. 4. Factorial experiments in the laboratory showed that Pardosa but not Grammonota reduced planthopper prey populations when prey density was low. By contrast, at high prey densities, both Pardosa and Grammonota had significant adverse effects on planthopper populations. Moreover, there was an interactive effect such that Grammonota reduced planthopper populations relatively more when Pardosa was absent than when it was present. 5. There was direct evidence for the intraguild predation of Grammonota by Pardosa such that fewer Grammonota survived in the presence of Pardosa than when it was absent. This result occurred whether planthopper prey were abundant or not. 6. Field releases of Grammonota in open plots resulted in significant but small decreases in the density of planthopper prey, both nymphs and adults. 7. Enhancing densities of Pardosa in open plots resulted in Grammonota suppression. The intraguild predation of Grammonota at this enhanced Pardosa density, however, did not preclude Pardosa from significantly reducing planthopper populations. 8. Although there was evidence that Grammonota reduced planthopper populations and that the intraguild predation of Grammonota by Pardosa occurred, the strength of these interactions was relatively weak given the low consumption rate of planthoppers by Grammonota (< 3 day^–1) and Grammonota by Pardosa (≈ 2 day^?1). Thus, weak asymmetric intraguild predation among spiders on the marsh likely dampens but does not eliminate the ability of Pardosa to exert significant top‐down control on planthopper populations.