Utilization, diet and diet selection by brent geese Branta bernicla bernicla on salt-marshes in Norfolk

The diet and utilization by brent geese of two plant communities ( Limonium/Armeria marsh—a short sward community in the upper mid salt-marsh and composed of several species, and Salicornia marsh—a community of the lower salt-marsh dominated by Salicornia europuea agg. and Aster tripolium ) were studied on the north Norfolk coast. The biomass on these communities was highest in autumn and declined markedly in November owing to the senescence and die-off of the succulent species. This decline was not significantly precipitated by the grazing of the geese but coincided with their switch to inland habitats where the biomass m^-2 and total food available on fields of winter wheat and grassland within the population's home range was 20–30 times greater. Brent geese continued to graze salt-marsh at a low intensity through the winter, feeding largely on Puccinellia maritima. In spring they returned largely to Limonium/Armeria marsh where they significantly reduced the regrowth of Puccinellia maritima.
The main plant species ealen was Puccinellia maritima , especially in mid winter. Salicornia maritima agg. leaves and seeds formed much of the diet in autumn, whereas Triglochin maritima, Plantago maritima and Aster tripolium were important foods on Limonium/Armeria marsh in spring. All these species were selected at some part of the year. Selection was related to the proline (an imino acid used by salt-marsh plants as an osmoregulator) and chloride content and avoidance related to tannins. Two species which contained condensed tannins. Limonium vulgare and Armeria maritima , were rarely eaten. As well as being a nutrient, proline may help to neutralize the effects of tannins in the diet if incorporated into proline-rich proteins.     

Molecular investigations in populations of Spartina anglica C.E. Hubbard (Poaceae) invading coastal Brittany (France)

Spartina anglica is a classical example of recent alloploid speciation. It arose during the end of the nineteenth century in England by hybridization between the indigenous Spartina maritima and the introduced East-American Spartina alterniflora. Duplication of the hybrid genome (Spartina x townsendii) gave rise to a vigorous allopolyploid involved in natural and artificial invasions on different continents. Spartina anglica was first recorded in France in 1906, and since then, it has spread all along the western French coast. Earlier studies revealed that native British populations display consistent morphological plasticity and lack of isozyme variation. In this paper, we use different molecular markers (randomly amplified polymorphic DNA, intersimple sequence repeats and restriction patterns from nuclear and chloroplast DNA sequences) to analyse the genetic patterns of the French populations of S. anglica. Our results show that French populations are mainly composed of one "major" multilocus genotype. This genotype is identical to the first-generation hybrid S. x townsendii from England. Losses of few markers from this genotype are observed but are restricted to a few populations from Brittany; it is likely that they appeared independently, subsequent to their introduction. In southern Brittany, no hybrids between S. anglica and S. maritima have been found where the two species co-occur. All French populations of S. anglica display the same chloroplast DNA sequences as S. alterniflora, the maternal genome donor. These findings are consistent with a severe genetic bottleneck at the time of the species formation, as a consequence of a unique origin of the species. Both parental nuclear sequences are present in the allopolyploid populations, revealing that for the markers investigated, no extensive changes have occurred in this young species.     

Origin and genetic diversity of Spartina anglica (Poaceae) using nuclear DNA markers

Spartina alterniflora, introduced into the UK in the 1800s, was the seed parent in an interspecific hybridization with S. maritima. The sterile F1 hybrid S. ×townsendii gave rise to the fertile allopolyploid S. anglica by chromosomal doubling. Previous chromosome, isozyme, and cpDNA surveys did not reveal notable genetic variation within either the parental or the hybrid species. We used nuclear DNA markers (random amplified polymorphic DNA ([RAPD]) and inter-simple sequence repeats (ISSR) to further explore the origin, diversity, and parentage of S. anglica. We found DNA fragments in S. ×townsendii were the aggregate of diagnostic DNA fragments from S. maritima and S. alterniflora, thus confirming its hybrid origin. The S. ×townsendii genotype was identical to most of the S. anglica individuals analyzed, establishing the genetic concordance of these two taxa. We found widespread genetic variation within S. anglica. This could indicate that S. anglica arose several times, from different S. maritima sires. Alternatively, alleles could have been lost through recombination and/or through loss of entire chromosomes in S. anglica. Finally, all but one S. anglica individual had a S. alterniflora component that was indistinguishable from a S. alterniflora plant extant in Marchwood, UK, leaving open the possibility that this plant is the actual seed parent of S. anglica.     

The evolution of Spartina anglica C.E. Hubbard (Gramineae): origin and genetic variability

An extensive survey of isozyme phenotypes in British populations of the amphidiploid salt marsh grass Spartina anglica and its putative parents has confirmed that the species arose by chromosome doubling in S. × townsendii , a sterile hybrid between S. maritima and S. alterniflora. Isozyme phenotypes and seed protein profiles indicate that S. anglica is almost totally lacking in genetic variation. Isozyme evidence also indicates that the parental species are characterized by low levels of genetic variation. The lack of variation in S. anglica is proposed as being due to a narrow genetic base resulting from a single origin, or a multiple origin from uniform parents; the fact that many populations are derived from very small founder populations; and because preferential pairing between identical homologous chromosomes prevents recombination between the divergent component genomes of the species. The low levels of isozyme variation that occur appear to be due to chromosome loss.
The consequences for the future evolution of S. anglica , given its lack of genetic variation, are discussed.     

Genetic and epigenetic consequences of recent hybridization and polyploidy in Spartina (Poaceae)

To study the consequences of hybridization and genome duplication on polyploid genome evolution and adaptation, we used independently formed hybrids (Spartina x townsendii and Spartina x neyrautii) that originated from natural crosses between Spartina alterniflora, an American introduced species, and the European native Spartina maritima. The hybrid from England, S. x townsendii, gave rise to the invasive allopolyploid, salt-marsh species, Spartina anglica. Recent studies indicated that allopolyploid speciation may be associated with rapid genetic and epigenetic changes. To assess this in Spartina, we performed AFLP (amplified fragment length polymorphism) and MSAP (methylation sensitive amplification polymorphism) on young hybrids and the allopolyploid. By comparing the subgenomes in the hybrids and the allopolyploid to the parental species, we inferred structural changes that arose repeatedly in the two independently formed hybrids. Surprisingly, 30% of the parental methylation patterns are altered in the hybrids and the allopolyploid. This high level of epigenetic regulation might explain the morphological plasticity of Spartina anglica and its larger ecological amplitude. Hybridization rather than genome doubling seems to have triggered most of the methylation changes observed in Spartina anglica.     

Molecular evidence for the maternal parentage in the hybrid origin of Spartina anglica C.E. Hubbard

Spartina anglica is a textbook example of a natural amphiploid, which originated from hybridization between S. alterniflora and S. maritima . Which of these species was the maternal parent has remained a mystery. Inheritance of chloroplast DNA in most angiosperms is strictly maternal and can thus be used to test the parentage of hybrid taxa. The DNA sequence of the chloroplast leucine tRNA gene intron was used to show that the introduced North American S. alterniflora is the female parent of the F ₁ hybrid S. x townsendii and the amphiploid S. anglica . A possible scenario for their origin is given.     

The evolution of Spartina anglica G. E. Hubbard (Gramineae): genetic variation and status of the parental species in Britain

The perennial salt marsh grass Spartina anglica is one of the classic examples of allopolyploid speciation. It originated on the south coast of England at the end of the nineteenth century following chromosome doubling in S. × townsmdii , a hybrid between the native British S. marilima and a species introduced from the United States, S. alterniflora. The nature of the origin of S. anglica is beyond doubt; however, it is not known whether it had a single or multiple origin. In order to address this problem we undertook a survey of the genetic variation in the parental species of S. anglica using isozyme electrophoresis. The results show that S. alterniflora has no detectable variation and that S. maritima has extremely low levels of variation. These results, unfortunately, prevent the question of a single or multiple origin from being answered. Possible reasons for the low levels of variation and its influence on the future of the species are discussed. Another problem concerning the parental species is the rapid decline of S. maritima in Britain. It is often assumed that the major factor in this regression is the invasion of its habitats by S. anglica. We have examined the status of S. marilima throughout its range in Britain and have found that S. anglica rarely co-occurs with S. maritima. We propose that the decline of S. maritima is largely due to the physical erosion of its habitats and that this erosion may produce suitable habitats for colonization by S. anglica.     

Retrotransposons and genomic stability in populations of the young allopolyploid species Spartina anglica C.E. Hubbard (Poaceae)

Spartina x townsendii arose during the end of the 19th century in England by hybridization between the indigenous Spartina maritima and the introduced Spartina alterniflora, native to the eastern seaboard of North America. Duplication of the hybrid genome gave rise to Spartina anglica, a vigorous allopolyploid involved in natural and artificial invasions on several continents. This system allows investigation of the early evolutionary changes that accompany stabilization of new allopolyploid species. Because allopolyploidy may be a genomic shock, eliciting retroelement insertional activity, we examined whether retrotransposons present in the parental species have been activated in the genome of S. anglica. For this purpose we used inter-retrotransposon amplified polymorphism (IRAP) and retrotransposons-microsatellite amplified polymorphism (REMAP) markers, which are multilocus PCR-based methods detecting retrotransposon integration events in the genome. IRAP and REMAP allowed the screening of insertional polymorphisms in populations of S. anglica. The populations are composed mainly of one major multilocus genotype, identical to the first-generation hybrid S. x townsendii. Few new integration sites were encountered in the young allopolyploid genome. We also found strict additivity of the parental subgenomes in the allopolyploid. Both these findings indicate that the genome of S. anglica has not undergone extensive changes since its formation. This contrasts with previous results from the literature, which report rapid structural changes in experimentally resynthesized allopolyploids.     

Spartina anglica C. E. Hubbard: a natural model system for analysing early evolutionary changes that affect allopolyploid genomes

Spartina anglica arose during the end of the 19th century in England by hybridization between the indigenous Spartina maritima and the introduced East American Spartina alterniflora and following genome duplication of the hybrid ( S.  ×  townsendii ). This system allows investigations of the early evolutionary changes that accompany stabilization of a new allopolyploid species in natural populations. Various molecular data indicate that S. anglica has resulted from a unique parental genotype. This young species contains two distinctly divergent homoeologous genomes that have not undergone extensive change since their reunion. No burst of retroelements has been encountered in the F₁ hybrid or in the allopolyploid, suggesting a 'structural genomic stasis' rather than 'rapid genomic changes'. However, modifications of the methylation patterns in the genomes of S.  ×  townsendii and S. anglica indicate that in this system, epigenetic changes have followed both hybridization and polyploidization.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 475–484.     

An Ecophysiological Comparison of Measurements of the Diurnal Rhythm of the Leaf Elongation and Changes of the Leaf Thickness of Salt-resistant Dicotyledonae and Monocotyledonae

Rozema, J., Arp, W., van Diggelen, J., Kok, E. and Letschert,J. 1987. An ecophysiological comparison of measurements of thediurnal rhythm of the leaf elongation and changes of the leafthickness of salt-resistant Dicotyledonae and Monocotyledonae.—J.exp. Bot. 38: 442–453. The continuous measurement of leaf elongation and leaf thicknesswith the use of a rotation potentiometer set up revealed a rapidand sensitive reaction of halophytic plants to conditions affectingthe plant's water relations. At increased salinity (450 molm^–3 NaCl) the rate of leaf elongation decreased both inAster tripolium and in Sparlina anghca. Increased shrinkageduring the day and a long period for recovery swelling at nightin leaves of Aster iripolium at increased salinity illustratesthat water shortage is part of the cause of salinity-inducedgrowth reduction. All dicotyledonous species analysed (Aster tripolium, A triplexhastata, A. littoralis, Suaeda maritima and Beta vulgaris) showeda day/night ratio of the leaf elongation rate lower than 1,while this ratio was higher than or equal to 1 in Monocotyledons(Spartina anglica, Juncus gerardii, J. maritimus, Festuca rubrassp. litoralis, Elymus pycnanthus). With the exception of Triglochinmaritima none of the monocotyledonous halophytes tested (Sparlinaanglica, Juncus gerardii, J. maritimus, Festuca rubra ssp. litoralis,Elymus pycnanthus) exhibited a diurnal rhythm of leaf thicknesschanges, such as was observed for all dicotyledonous speciesstudied (Aster tripolium, Atriplex hastata, A. littoralis, Salicorniabrachyslachya, Suaeda maritima, Glaux maritima, Odontites vernassp. serotina). The diurnal pattern of the leaf elongation rateand the leaf thickness changes can be explained by variationof photosynthetic rate and transpiration water losses by stomatalclosure in the dark and opening in the light such as shown forthe dicotyledon species Glaux maritima. This difference betweendicot and monocot species in diurnal variation of the leaf elongationrate and leaf thickness may partly be explained in terms ofthe different position of the growth zone and possibly by adifference in elasticity of the tissue of halophytic monocotyledonsand dicotyledons. The consequences of these differences arediscussed. Key words: Leaf elongation rate, leaf thickness, water relations, salt resistance, Dicotyledonae, Monocotyledonae     

米草属引入中国海岸带的利弊分析

1963年以来,中国先后从国外引进3种米草。实践证明,大米草和互花米草是盐渍土壤植物群落形成的先锋植物,适应于中国海岸带自然环境,生长发育正常,具有明显的促淤造陆和保护海岸等生态功能,并具有显著的经济效益.大米草和互花米草在南方海岸繁殖过快是近岸水体富营养化的结果,但并未对环境、动植物种群和人类构成侵害,其在北方海岸未发生异常现象,故应客观地评估米草的功与过。本文从利弊2个方面对中国海岸滩涂引入米草进行了分析。     

A comparative study of the tolerance of salt marsh plants to manganese

Summary Salicornia europaea, Puccinellia maritima, Triglochin maritima, Aster tripolium, Plantago maritima, Armeria maritima, Juncus gerardii andFestuca rubra, collected as seed from a salt marsh at Portaferry, County Down, were grown on saline (340 mM NaCl) and non saline nutrient solutions at five concentrations of manganese sulphate (0.025–10.0 mM). After an eight week growing period, shoot and root yields and the concentrations of sodium, potassium, calcium and manganese in the shoots were determined.Except forS. europaea the saline treatments had a strongly limiting effect on plant growth. Each of the species investigated showed a degree of tolerance to high concentrations of manganese which was similar to that of calcifuge species and plants characteristic of waterlogged sand dune slack communities, but which was very much greater than that ofArrhenatherum elatius a species usually excluded from acidic soils. There was little evidence to support the hypothesis that tolerance of high manganese concentrations was correlated with the position of the experimental plants in the salt marsh ecotone or that the manganese nutrition of halophytic and glycophytic marsh species differs. Whilst manganese uptake increased proportionally with solution manganese concentration, there were few other major effects of manganese on the balance of shoot cation concentrations in the plants investigated. Both antagonistic and synergistic effects of sodium on manganese uptake were recorded for different species.     

Molecular phylogeny of hybridizing species from the genus Spartina Schreb. (Poaceae)

Interspecific hybridization events have been reported in the genus Spartina Schreb. (Poaceae), involving the east American species Spartina alterniflora, and including either introgression (e.g., with the western American Spartina foliosa) or allopolyploid speciation (e.g., with the Euro-African Spartina maritima). Molecular phylogenetic analysis of the genus has been undertaken in order to understand phylogenetic relationships and genetic divergence among these hybridizing species. Twelve Spartina species have been sequenced for two nuclear DNA regions (ITS of ribosomal DNA, and part of the Waxy gene) and one chloroplast DNA spacer (trnT-trnL). Separate and conditional combined phylogenetic analyses using Cynodon dactylon as the outgroup have been conducted. Spartina is composed of two lineages. The first clade includes all hexaploid species: the Euro-African S. maritima (2n = 60), the East-American S. alterniflora (2n = 62) and the West-American S. foliosa (2n = 60). Spartina alterniflora appears as a closely related sister species to S. foliosa. Although belonging to the same lineage, Spartina maritima appears consistently more genetically differentiated from S. alterniflora than S. foliosa. The tetraploid species S. argentinensis (2n = 40) is placed at the base of this first clade according to the Waxy data, but its position is not well resolved by the other sequences. The second well-supported main lineage within genus Spartina includes the other tetraploid American species. Significant incongruence has been encountered between the waxy based tree and both the ITS and trnT-trnL trees concerning the position of S. densiflora, suggesting a possible reticulate evolution for this species. The results agree with hybridization patterns occurring in Spartina: introgression involving closely related species (S. alterniflora and S. foliosa) on one hand, and alloploid speciation involving more differentiated species (S. alterniflora and S. maritima) on the other hand.     

Molecular phylogenetics of Limonium and related genera (Plumbaginaceae): biogeographical and systematic implications

Phylogenetic relationships within Limonium (Plumbaginaceae) are evaluated using sequence data from three plastid regions (rbcL, the trnL intron, and the trnL-trnF intergenic spacer). Sixty-six species representing the major genera of Staticoideae, including representatives of all sections and genera formerly included in Limonium, have been analyzed using four species of Plumbaginoideae as an outgroup. Analyses of each separate and combined data set yield similar results. Afrolimon is embedded in Limonium and related to L. vulgare, the type of Limonium. Limonium is split into two major clades corresponding to subgenera, but otherwise the current infrageneric classification proved to be artificial. Some groups restricted to particular areas can be recognized, and their synapomorphies are discussed. The presence of an isolated taxon in the Canary Islands is used as a calibration point for age estimates of the major events in the genus, including migrations to the Southern Hemisphere, the Canary Islands, and Asia. The rapid radiation of Limonium in the Mediterranean basin appears to coincide with the desiccation of the Mediterranean Sea in the Messinian (late Miocene).     

An in Vivo Nuclear Magnetic Resonance Investigation of Ion Transport in Maize (Zea mays) and Spartina anglica Roots during Exposure to High Salt Concentrations

The response of maize (Zea mays L.) and Spartina anglica root tips to exposure to sodium chloride concentrations in the range 0 to 500 mM was investigated using 23Na and 31P nuclear magnetic resonance spectroscopy (NMR). Changes in the chemical shift of the pH-dependent 31P-NMR signals from the cytoplasmic and vacuolar orthophosphate pools were correlated with the uptake of sodium, and after allowing for a number of complicating factors we concluded that these chemical shift changes indicated the occurrence of a small cytoplasmic alkalinization (0.1-0.2 pH units) and a larger vacuolar alkalinization (0.6 pH units) in maize root tips exposed to salt concentrations greater than 200 mM. The data were interpreted in terms of the ion transport processes that may be important during salt stress, and we concluded that the vacuolar alkalinization provided evidence for the operation of a tonoplast Na+/H+-antiport with an activity that exceeded the activity of the tonoplast H+ pumps. The intracellular pH values stabilized during prolonged treatment with high salt concentrations, and this observation was linked to the recent demonstration (Y. Nakamura, K. Kasamo, N. Shimosato, M. Sakata, E. Ohta [1992] Plant Cell Physiol 33: 139-149) of the salt-induced activation of the tonoplast H+- ATPase. Sodium vanadate, an inhibitor of the plasmalemma H+- ATPase, stimulated the net uptake of sodium by maize root tips, and this was interpreted in terms of a reduction in active sodium efflux from the tissue. S. anglica root tips accumulated sodium more slowly than did maize, with no change in cytoplasmic pH and a relatively small change (0.3 pH units) in vacuolar pH, and it appears that salt tolerance in Spartina is based in part on its ability to prevent the net influx of sodium chloride.     

Changes in a salt-marsh vegetation as a result of grazing and mowing — A five-year study of permanent plots

Summary Part of a salt-marsh (32 ha), ungrazed from 1958 until 1971, was grazed again from 1972 onwards with young cattle (1.3–1.7 per ha, May–October). In five vegetation types management experiments, including doing nothing (control). June mowing, August mowing, June and August mowing, all in combination or not in combination with grazing have been started with the objective to compare annually their effects on the vegetational structure and composition by means of permanent plots (2×2 m).Thirteen years (1958–1971) after the grazing had ceased the vegetation types of Festuca rubra/Armeria maritima, Artemisia maritima, Juncus maritimus and Elytrigia pungens hardly changed anymore. The vegetation type of Festuca rubra/Limonium vulgare, changed considerably. The experiments showed rather specific effects during the period 1971–1975 for each type of vegetation. Changes in the Festuca rubra/Armeria maritima vegetation were small and gradually under all treatments. Changes in The Artemisia maritima, Festuca rubra/Limonium vulgare and Juncus maritimus vegetation types were rather great under the different grazing treatments, the changes being abrupt and especially taking place in the third and fourth year of the experiments. The Elytrigia pungens vegetation showed large changes under all treatments, except the control plot, whereas these changes were abrupt, particularly occurring in the first and second year of the experiments. The study is continued.Nomenclature of taxa follows Heukels & van Ooststroom (1970), that of syntaxa Westhoff & den Held (1969).Contribution to the Symposium of the Working Group for Succession research on permanent plots of the International Society for Vegetation Science, held at Yerseke, the Netherlands, October 1975.Thanks are due to Prof. Dr. D. Bakker for his interest in the present study and his critical reading of the text, and to Drs D.C.P. Thalen for critical reading and correction of the English text. Thanks are also due to the students who analyzed the permanent plots: M. van der Duim and F. Prins (1971), M. van der Duim (1972), R. Schwab and T. Schwab-Vos (1973), and G. Allersma (1974, 1975).     

Uniformity of the nuclear and chloroplast genomes of Spartina maritima (Poaceae), a salt-marsh species in decline along the Western European Coast

Spartina maritima is a salt-marsh species from European and African Atlantic coasts. In the northern range of the species (including north-west France), a rapid decline of the populations has been observed during the 20th century. In this paper, the molecular diversity of 10 populations of S. maritima from France has been investigated using nuclear and chloroplast DNA markers: inter-simple sequence polymorphism (ISSR), randomly amplified polymorphic DNA (RAPD), inter-retrotransposon amplified polymorphism (IRAP), and PCR-RFLP of a 5 kb long portion of chloroplast DNA. The results reveal an extremely low level of genetic variation in this species: only one nuclear marker (out of 98) was polymorphic, with the presence of two genotypes randomly distributed among the populations. The lack of genetic diversity is interpreted in light of the almost exclusive vegetative propagation of the species in its northern range, the colonization history of the populations, and the origin of S. maritima (2n = 60) in the hexaploid lineage of the genus and in the context of the management of S. maritima populations.     

Nitrogen and phosphorus limitation in a coastal barrier salt marsh: the implications for vegetation succession

1 A factorial fertilizer experiment was conducted in a 15-year-old coastal barrier salt marsh with a low soil nitrogen content, and in an older 100-year-old marsh with a higher nitrogen content. Plots were fertilized at high and low marsh elevations in both marshes. Nitrogen and phosphorus were applied at low and high concentrations both separately and in combination in each of 3 successive years.
2 Nitrogen limited above-ground plant growth in both young and old salt marshes in all years. Phosphorus limitation of plant growth was apparent in the first year in the young marsh and in the last year in both marshes. In young marshes with low soil organic matter, phosphorus limitation may occur. In addition, phosphorus limitation occurs at both successional stages when a marsh is saturated with nitrogen.
3 Plant species that are typical of nitrogen-rich habitats and late successional stages significantly increased in biomass after fertilization. Limonium vulgare , a low stature species of early and intermediate successional stages, decreased in biomass, whereas the taller Elymus pycnanthus and Artemisia maritima increased. After 3 years of fertilization, plant species composition in a young marsh was similar to the species composition in an unfertilized older marsh. Fertilization of a 100-year-old marsh, however, still resulted in a change in plant species composition, suggesting that succession was still occurring and that, overall, plants in marshes of different age are similar in their response to fertilization.     

初始克隆分株数对大米草表型可塑性及生物量分配的影响

克隆植物大米草 (Spartina anglica) 目前在我国出现了严重的自然衰退 (Dieback),为了阐明大米草衰退的机理,分析影响大米草形态可塑性的因素与自然衰退之间的相关性,以期为近缘植物互花米草 (S. alterniflora) 这一爆发种群的生物控制提供借鉴,对3种不同初始克隆分株数 (单克隆、三克隆和五克隆) 大米草的克隆生长、生物量累积与分配和异速生长特征进行了野外栽培试验。研究结果表明,初始克隆分株数对间隔子长度影响较弱;初始多克隆的分支强度高于初始单克隆;初始三克隆和五克隆在总生物量 (7.921 5~10.431 7 g 和 8.903 9~10.431 7 g)、地上生物量 (3.396 1~4.255 8 g 和3.618 4~4.338 9 g)、地下生物量 (4.286 9~5.206 6 g 和 5.298 8~6.079 3 g)和根状茎生物量 (1.318 6~1.767 7 g 和 1.499 1~2.038 7 g) 积累上均显著高于初始单克隆,不同初始克隆分株数条件下根生物量差异不显著;初始多克隆倾向于将资源更多地分配给根状茎,而初始单克隆倾向于将更多的资源分配给根系。由此推断,在不同初始克隆分株数条件下,大米草的形态可塑性和生物量分配格局的差异显示出在同样资源格局下,初始多克隆的克隆生殖能力较初始单克隆强。初始多克隆生长的大米草较初始单克隆生长的大米草更能占据优势生境,选择生境“觅养”的能力与克隆繁殖能力更强。     

Nitrogen level changes the interactions between a native (Scirpus triqueter) and an exotic species (Spartina anglica) in Coastal China

The exotic species Spartina anglica, introduced from Europe in 1963, has been experiencing a decline in the past decade in coastal China, but the reasons for the decline are still not clear. It is hypothesized that competition with the native species Scirpus triqueter may have played an important role in the decline due to niche overlap in the field. We measured biomass, leaf number and area, asexual reproduction and relative neighborhood effect (RNE) of the two species in both monoculture and mixture under three nitrogen levels (control, low and high). S. anglica showed significantly lower biomass accumulation, leaf number and asexual reproduction in mixture than in monoculture. The inter- and intra-specific RNE of S. anglica were all positive, and the inter-specific RNE was significantly higher than the intra-specific RNE in the control. For S. triqueter, inter- and intra-specific RNE were negative at the high nitrogen level but positive in the control and at the low nitrogen level. This indicates that S. triqueter exerted an asymmetric competitive advantage over S. anglica in the control and low nitrogen conditions; however, S. anglica facilitated growth of S. triqueter in high nitrogen conditions. Nitrogen level changed the interactions between the two species because S. triqueter better tolerated low nitrogen. Since S. anglica is increasingly confined to upper, more nitrogen-limited marsh areas in coastal China, increased competition from S. triqueter may help explain its decline.