Geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata

We investigated geographic patterns of herbivory and resource allocation to defense, growth, and reproduction in an invasive biennial, Alliaria petiolata, to test the hypothesis that escape from herbivory in invasive species permits enhanced growth and lower production of defensive chemicals. We quantified herbivore damage, concentrations of sinigrin, and growth and reproduction inside and outside herbivore exclusion treatments, in field populations in the native and invasive ranges. As predicted, unmanipulated plants in the native range (Hungary, Europe) experienced greater herbivore damage than plants in the introduced range (Massachusetts and Connecticut, USA), providing evidence for enemy release, particularly in the first year of growth. Nevertheless, European populations had consistently larger individuals than US populations (rosettes were, for example, eightfold larger) and also had greater reproductive output, but US plants produced larger seeds at a given plant height. Moreover, flowering plants showed significant differences in concentrations of sinigrin in the invasive versus native range, although the direction of the difference was variable, suggesting the influence of environmental effects. Overall, we observed less herbivory, but not increased growth or decreased defense in the invasive range. Geographical differences in performance and leaf chemistry appear to be due to variation in the environment, which could have masked evolved differences in allocation.     

除草剂对黄河三角洲入侵植物互花米草的影响

互花米草(Spartina alterniflora)作为我国危害最严重的外来入侵植物之一,严重威胁滨海湿地生态系统安全。筛选可高效灭除互花米草的除草剂,为互花米草防治提供技术支持。2017年7月在黄河三角洲潮间带对互花米草茎叶喷施不同除草剂,在施药当年和次年对互花米草生长状况和大型底栖动物密度进行跟踪调查。研究结果表明:1)高效氟吡甲禾灵对互花米草的灭除效果最好,既能杀死互花米草地上部分从而完全抑制有性繁殖,又可完全抑制次年的无性繁殖;2)氰氟草酯对互花米草的灭除效果也比较好,可以完全抑制互花米草的生长和结穗,但抑制根状茎无性繁殖能力的效果稍差,草甘膦可以完全抑制互花米草的有性繁殖,但无法抑制次年互花米草的无性繁殖;3)施用除草剂在短期内会毒害某些底栖动物,但在1年后底栖动物种群数量与对照处理无显著差异。除草剂对环境的影响程度与除草剂用量和施用时间息息相关,未来研究中,应在保证灭草效果的前提下,探索最佳用药时间和最低用量以最大限度地降低环境影响。     

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.     

Patterns of resource allocation in male and female plants of Rumex acetosa and R. acetosella

Summary Patterns of resource allocation in the dioecious Rumex acetosa and R. acetosella were investigated. Males were found to allocate more to reproduction during flower production than females, whereas females invested considerably more in reproduction during seed production. Altogether, females allocated both a higher total amount and a higher proportion of energy to reproduction than did males. By regression analysis, the influence of plant size on reproductive effort was examined separately for males and females. The results indicated that while reproductive effort is sometimes lower for tall plants than for small plants, size-independent effects have a greater influence on reproductive effort than size distribution. An analysis of variance was conducted to investigate the effects of population, season, sex and their interactions on plant size, and an analysis of covariance was used to study differences in resource allocation patterns. Different interaction effects were found to be most important in the two species of Rumex.     

Competition between <Emphasis Type="Italic">Lythrum salicaria</Emphasis> and a rare species: combining evidence from experiments and long-term monitoring

The rare endemic plant Sidalcea hendersonii (Henderson’s checker-mallow) occurs in tidal marshes of the Pacific Northwest and may be threatened by Lythrum salicaria (purple loosestrife), a European invader plant. We compared the abundances of Lythrum and Sidalcea in a wetland in British Columbia (Canada) in 1999 to those measured in 1979 to track changes in both species. Although the frequency of Sidalcea decreased by more than 50%, and that of Lythrum increased by almost 20%, there was no significant relationship between the changes of the two species. We assessed the potential effects of competition by Lythrum on Sidalcea in field and patio experiments. In the field, we measured the response of Sidalcea to the removal of Lythrum over a two-year period and compared this to the response of Sidalcea to the removal of native species and in unmanipulated control plots. Removal of Lythrum significantly improved the vegetative performance of Sidalcea compared to the removal of randomly selected native plants and the control treatment in the first year. In the second year, the performance of Sidealcea did not differ significantly with treatment. Removals did not influence the reproductive performance of Sidalcea in either year. A one-year additive experiment, carried out in pots, compared the competitive effect of Lythrum on Sidalcea with that of two native species. Lythrum’s impact on Sidalcea was not consistently stronger than that of the native species. Collectively, these results do not indicate a strong impact of Lythrum on the reproduction or abundance of Sidalcea.     

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.     

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.     

Interactions between the grass shrimp <Emphasis Type="Italic">Palaemonetes pugio</Emphasis> and the salt marsh grasses <Emphasis Type="Italic">Phragmites australis</Emphasis> and <Emphasis Type="Italic">Spartina alterniflora</Emphasis>

The grass shrimp Palaemonetes pugio, a species common to Spartina alterniflora-dominated marshes, may be sensitive to the invasion of the common reed Phragmites australis in northeastern US salt marshes. We examined two questions: (1) Do grass shrimp have a preference for the native plant over the non-native plant? (2) Are grass shrimp more effective foragers on P. australis? We tested the first hypothesis by comparing the amount of time shrimp spend in physical contact with the plant types over a 1-h period. Shrimp were observed under different arrangements of vegetation to control for differences in conspicuous structural features. Additionally, the amount of time shrimp spent foraging on S. alterniflora and P. australis shoots was compared to determine if shrimp graze more often on S. alterniflora. Shrimp spent significantly more time in contact with S. alterniflora only when plant types were grouped at opposite ends of aquaria, but did not exhibit a foraging preference for this plant type. To address our second question, we investigated the effects of shrimp foraging on stem epifauna, an assemblage of semi-aquatic invertebrates associated with macrophyte shoots. Previous research showed that P. australis supports a lower density of stem-dwelling epifauna relative to S. alterniflora. We hypothesized that the primary grazer of this community, P. pugio, can forage on P. australis stems more effectively due to structural differences between the two plants, causing the lower abundance of epifauna through top-down effects. We exposed individual shoots inhabited by epifauna to shrimp and compared faunal densities on exposed shoots to densities on control shoots after 18 h. The reduction of epifauna by predation was proportional on the two plant types. Therefore, top-down effects can be ruled out as an explanation for the patchy distribution of epifauna observed in P. australis–S. alterniflora marshes.     

互花米草入侵影响下闽江河口湿地土壤有效硅的时空变化特征

2016年1—12月,选择闽江河口鳝鱼滩的短叶茳芏湿地、互花米草湿地以及二者的交错带湿地为研究对象,采用定位研究方法探讨了互花米草入侵影响下湿地土壤有效硅含量的时空变化特征。结果表明:互花米草入侵影响下3块湿地土壤有效硅含量随时间推移整体呈波动上升趋势;互花米草入侵显著提高了鳝鱼滩湿地30—60 cm土层土壤有效硅含量(P0.01),与短叶茳芏湿地相比,交错带湿地和互花米草湿地30—60 cm土层土壤有效硅含量分别增加了8.56%和19.97%,逐步线性回归分析表明土温和电导是影响其变化的重要因素(P0.01)。研究互花米草入侵影响下湿地土壤有效硅含量的变化特征,对于揭示湿地生态系统生源要素硅生物地球化学循环过程以及互花米草入侵及其扩张机制具有重要意义。     

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.     

Blocking Phragmites australis reinvasion of restored marshes using plants selected from wild populations and tissue culture

This study tested a vegetation strategy for controlling Phragmites australis invasion into brackish marshes as an alternative to the current technique of repeated herbicide sprays followed by burning. This strategy involves blocking P. australis by planting desired plants selected from wild populations and/or tissue culture regenerants at key points on the major routes of P. australis invasion. The planting of native species was conducted at three sites in a herbicide-treated P. australis marsh near Salem, NJ. Wild population selections of three upland marsh shrubs, Myrica cerifera, Baccharis halimifolia, and Iva frutescens, as well as two grass species, Spartina alterniflora and Spartina patens, and two rushes, Juncus gerardi and Juncus roemerianus, were planted according to their normal zonation positions. Tissue culture regenerated plants of the two grasses and two rushes, and the sedge species Scirpus robustus, were also planted. Plant growth at each site was monitored each year after planting for up to 3 years. Most plants of B. halimifolia, I. frutescens, J. roemerianus, and S. patens demonstrated a consistent vigorous growth at all three sites, whether or not the plants were collected from wild populations or were tissue culture regenerants. These multi-layered walls of plants demonstrated effectiveness in controlling the P. australis by restricting or inhibiting its spread. Upon screening 48 regenerated plants of S. patens at one of the three sites, we found that some regenerants showed enhanced characteristics for blocking P. australis, such as greater expansion and a high stem density. The availability of the tissue culture-regenerated plants of the native marsh species makes it possible to select lines from local genotypes that have desirable characteristics for wetland restoration projects, such as blocking P. australis reinvasion.     

Population demographics and trade-offs to reproduction of an invasive and noninvasive species of <Emphasis Type="Italic">Rubus</Emphasis>

Do trade-offs between growth and reproduction differ between an invasive and noninvasive plant species and how do such trade-offs relate to population demographics? To help address these questions, we compared demographics for an invasive plant species, Rubus discolor, with a noninvasive congener, R. ursinus, in several populations of varying density. Removal of floral buds from reproductive canes increased the size of juvenile canes that arose from clonal sprouting in R. ursinus, suggesting a trade-off between current reproduction and growth. Removal of floral buds had no effect on growth of R. discolor. R. ursinus displayed trade-offs between reproduction (sexual and vegetative) and future growth based on negative correlations between leaf area production and both clonal sprouting and seedling production during the previous year. R. discolor did not exhibit these trade-offs. Both species had high population growth rates in low-density populations, but exhibited little or no growth in high-density populations. A life table response experiment was used to determine the underlying cause for the effect of density on population growth. For R. ursinus, lack of population growth in high-density populations was due primarily to increased mortality of clonally sprouting canes, while for R. discolor, it was due to decreased clonal cane production. Elasticity analysis revealed that clonal growth was more important than sexual reproduction for population growth of both species. However, elasticity values for sexual reproduction in R. discolor were greater in high- than low-density populations. This suggests an increased reliance on sexual reproduction in populations that had reached stable sizes, which could increase the capacity of R. discolor to disperse to new sites. Elasticity analyses were also used to simulate the efficacy of various control strategies for R. discolor. Control of this species could be attained by reducing clonal production within existing populations while reducing seed production to limit establishment of new populations.     

Lacunal allocation and gas transport capacity in the salt marsh grass Spartina alterniflora

Summary Lacunal allocation as the fraction of the total cross sectional area of leaves, stem bases, rhizomes, and roots was determined in both tall and short growth forms of Spartina alterniflora collected from natural monospecific stands. The results indicate that in both growth forms lacunal allocation is greater in stem bases and rhizomes than in leaves and roots and that tall form plants allocate more of their stem and rhizome to lacunae than short form plants.Measurements made in natural stands of Spartina alterniflora suggest that total lacunal area of the stem base increases with increasing stem diameter and that stem diameter increases with increasing plant height and above-ground biomass. However, the fraction of cross section allocated to lacunae was relatively constant and increased only with the formation of a central lacuna.Experimental manipulations of surface and subsurface water exchange were carried out to test the influence of flooding regime on aerenchyma formation. No significant differences in lacunal allocation were detected between plants grown in flooded (reduced) and drained (oxidized) sediments in either laboratory or field experiments. While aerenchyma formation in Spartina alterniflora may be an adaptation to soil waterlogging/anoxia, our results suggest that lacunal formation is maximized as a normal part of development with allocation constrained structurally by the size of plants in highly organic New England and Mid-Atlantic marshes.The cross sectional area of aerenchyma for gas transport was found to be related to the growth of Spartina alterniflora with stands of short form Spartina alterniflora exhibiting a lower specific gas transport capacity (lacunal area per unit below ground biomass) than tall form plants despite having a similar below-ground biomass supported by a 10 fold higher culm density. The increased specific gas transport capacity in tall vs. short plants may provide a new mechanism to explain the better aeration, higher nutrient uptake rates and lower frequency of anaerobic respiration in roots of tall vs. short Spartina alterniflora.     

Effects of growing conditions on the growth of and interactions between salt marsh plants: implications for invasibility of habitats

A common but often less tested explanation for the successful invasion of alien species is that invasive alien species outcompete their co-occurring natives, which may not always be the case. In this study, we established artificial environmental gradients in a series of pot experiments with controlled environments to investigate the effects of salinity, sediment type and waterlogging on the performance of and interactions between Phragmites australis (native) and Spartina alterniflora (alien), which generally co-exist in the saline intertidal zones of Chinese and American coasts. Significant effects of salinity and waterlogging were detected on biomass production and morphological characteristics of S. alterniflora and P. australis, and the competitive interactions between the two species were found to vary with all three environmental factors in our experiments. Relative Neighbor Effect (RNE) analyses indicate that competitive dominance of S. alterniflora occurred under the conditions of high salinity, sandy sediment and full immersion, whereas P. australis showed competitive dominance under the conditions of low salinity and non-immersion. Our results suggest that S. alterniflora might outcompete P. australis under conditions present in early salt marsh succession, which support the viewpoint that the outcomes of competition between co-occurring native and invasive alien plants depend on the growing conditions. The implication of this study is that in response to the environmental changes expected from seawater intrusion and sea-level rise, the range of S. alterniflora is expected to expand further in the Yangtze River estuary in the future.     

A Panax-centric view of invasive species

Plant-centric sampling provides a novel approach to quantifying the potential impact of invasive species on native plant species. The aim of this study was to determine the level of exposure of individuals and populations of Panax quinquefolius to invasive plant species using this approach in thirty natural ginseng populations. A high level of invasion was found with 63–70% of ginseng populations containing at least one invasive species. Approximately one-third of all individuals were found in close proximity to invasive plants. The most prevalent invasive species were Rosa multiflora and Berberis thunbergii. The exposure to invasives of plants in different size classes varied among populations. Invasive species presence increased with greater ginseng population sizes and presence of harvest. The abundance of invasives plants within forest interiors near this valuable medicinal herb suggests that the economic and ecological costs of competitive interactions with native species could be high.     

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.     

Distinct life histories of peracarid crustaceans in a Ria Formosa salt marsh (S. Portugal)

Pitfall traps were exposed between February 1993 andSeptember 1994 every 4th to 6th week near the highwater mark in a Ria Formosa salt marsh (Portugal).With only a few exceptions, peracarid crustaceansaccounted for more than 95% of the individualscaught. Three amphipod species (Orchestiagammarellus(Pallas), Orchestia mediterraneaCosta, Talorchestia deshayesii(Audouin)) and3 isopod species (Tylos ponticus(Arcangeli),Halophiloscia couchi(Kinahan) and Porcellio lamellatus(Uljanin) Budde-Lund) wereidentified. In general, these species were most activefrom spring to autumn.Data collected included information on growth rate,life expectancy and timing of reproduction. Theamphipods displayed more opportunistic life-historypatterns with high growth rates, reproductive activityduring most of the year, early sexual maturity andrelatively short life expectancies. The isopods weremore persistent with slower growth rates, morerestricted reproductive periods and longer lifeexpectancies.There is no indication in the literature that most ofthese species are particularly common in salt marshes.In addition, peracarid crustaceans are not mentionedas dominant taxa in most studies of salt marsh fauna.We suggest that the peracarids play an important rolein the degradation of organic matter in salt marshesand hypothesize that the high numbers of peracaridsfound in Ria Formosa is due to the high contributionof leaf shedding plants.     

Mast-seeding in the cycad genus Encephalartos: a test of the predator satiation hypothesis

Mast-seeding behaviour was monitored in 18 populations of eight species of the African cycad genus Encephalartos between 1988 and 1991. The coefficient of variation (V) in annual cone production for each population ranged between 88 and 200, indicating large fluctuations in reproductive effort between years. Data were collected to determine whether mast-seeding reduced levels of predispersal seed predation by satiating seed predators in mast years and whether it resulted in a reproductive advantage over plants which reproduced more frequently. Masting intensity was greatest in those populations in which individual plants suffered the highest levels of predispersal seed predation in years when only a few plants produced seeds. The principal seed predators were two congeneric weevil species, Antliarhinus zamiae and A. signatus, which develop exclusively on cycad seeds. The lowest intensity of mast-seeding was recorded for cycad populations with low levels of seed predation and in which A. zamiae and A. signatus occurred only in low numbers or were entirely absent. Larger seed crops appeared to result in lower levels of seed predation by A. zamiae and A. signatus in four populations of E. altensteinii, and differences in seed crop size accounted for 48–66% of variation in levels of seed predation in populations of five cycad species. In one population of E. altensteinii, lower levels of seed predation in plants reproducing periodically resulted in a reproductive advantage over plants reproducing more frequently. These results are consistent with the predator satiation hypothesis. However, in most cycad populations, numbers of seed predators did not appear to decrease significantly after a period of 2–8 years between reproductive episodes and, in two of three populations examined, periodic reproduction did not increase the number of seeds surviving to dispersal over a 4-year period. These results are interpreted to mean that periodic reproduction has not evolved in response to selection imposed by seed predators, but that selection may favour those plants which experience lower levels of seed predation by coning in synchrony with the majority of plants in the population.     

Utilization of a citizen monitoring protocol to assess the structure and function of natural and stabilized fringing salt marshes in North Carolina

Narrow fringing salt marshes dominated by Spartina alterniflora occur naturally along estuarine shorelines and provide many of the same ecological functions as more extensive marshes. These fringing salt marshes are sometimes incorporated into shoreline stabilization efforts. We obtained data on elevation, salinity, sediment characteristics, vegetation and fish utilization at three study sites containing both natural fringing marshes and nearby restored marshes located landward of a stone sill constructed for shoreline stabilization. During the study, sediment accretion rates in the restored marshes were approximately 1.5- to 2-fold greater than those recorded in the natural marshes. Natural fringing marsh sediments were predominantly sandy with a mean organic matter content ranging between 1.5 and 6.0%. Average S. alterniflora stem density in natural marshes ranged between 130 and 222 stems m⁻², while mean maximum stem height exceeded 64 cm. After 3 years, one of the three restored marshes (NCMM) achieved S. alterniflora stem densities equivalent to that of the natural fringing marshes, while percentage cover and maximum stem heights were significantly greater in the natural than in the restored marshes at all sites. There was no significant difference in the mean number of fish, crabs or shrimp captured with fyke nets between the natural and restored marshes, and only the abundance of Palaemonetes vulgaris (grass shrimp) was significantly greater in the natural marshes than in the restored ones. Mean numbers of fish caught per 5 m of marsh front were similar to those reported in the literature from marshes adjacent to tidal creeks and channels, and ranged between 509 and 634 fish net⁻¹. Most of the field data and some of the sample analyses were obtained by volunteers as they contributed 223 h of the total 300 h spent collecting data from three sites in one season. The use of fyke nets required twice as many man-hours as any other single task. Vegetation and sediment parameters were sensitive indicators of marsh restoration success, and volunteers were capable of contributing a significant portion of the labor needed to collect these parameters. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

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.