长江口九段沙湿地不同生境中大型底栖动物群落结构特征分析

2004年10月至2005年1月对九段沙湿地大型底栖动物的群落分布特征及环境因子进行了调查。选取6种主要生境的57个样方,分析大型底栖动物的种类、密度、生物量、多样性指数及均匀度指数等群落特征。共记录到28种大型底栖动物,主要由甲壳动物、环节动物、软体动物及昆虫幼体组成,优势种为谭氏泥蟹(Ilyrplaxdeschampsi)、光滑狭口螺(Stenothyraglabra)、堇拟沼螺(Assimineaviolacea)、绯拟沼螺(A.latericea)和霍甫水丝蚓(Limnodriuhoffmeisteri)。6种生境类型中以低潮位光滩大型底栖动物种类最为丰富,但其密度最低;海三棱藨草(Scirpusmariqueter)带底栖动物密度最高,并且其均匀度与其他生境存在显著差异;互花米草(Spartinaalterniflora)带下底栖动物的种类组成与其他生境有一定差异,但种类数、密度和多样性处于平均水平,总体群落结构没有显著差异。     

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.     

九段沙湿地植被群落演替与格局变化趋势

长江口新生湿地九段沙由上沙、中沙和下沙三部分组成。2004年对湿地植被群落的演替与格局变化进行了研究。结果表明,九段沙湿地植被群落由芦苇(Phragmites austra-lis)、互花米草(Spartina alterniflora)、海三棱藨草(Scirpus mariqueter)和藨草(S.triqueter)构成。芦苇和互花米草生长于潮滩高程,海三棱藨草和藨草生长于低潮位。1998—2004年,上沙植被覆盖面积年平均约增长17%,中沙22%,下沙38%~39%,植被群落处于快速增长演替状态,这可能与沙洲发育速度有关。由于在中沙、下沙人为种植过外来物种互花米草,其增长速度最快,覆盖面积占九段沙植被总面积的比例从1998年不足1%上升到2004年的22%,地上部分总生物量的年平均增长率为79.79%,互花米草对藨草-海三棱藨草的竞争压力较大。鉴于互花米草对九段沙的影响尚无定论,本文提出了初步的植被管理措施。     

Ecological differentiation and reproductive isolation of two closely related sympatric species of Oenanthe (Apiaceae)

Oenanthe conioides is a lower Elbe endemic plant species growing in the freshwater tidal zone around Hamburg (Germany). Its closest relative Oenanthe aquatica is widely distributed in Eurasia and grows in calm and shallow freshwater. The two species differ in habitat requirements but are otherwise sympatrically distributed, suggesting that ecological divergence has to be maintained in the face of gene flow. In the present study, we investigated ecological differentiation and reproductive isolation in these two species. An amplified fragment length polymorphism analysis found clear genetic differentiation between the two species implying reproductive isolation. A reciprocal transplantation experiment including artificial F₁ hybrids showed strong selection against immigrants. In the two parental habitats, the non‐native species are less fit than the native species. Hybrids are less fit in the habitat of Oe. aquatica but perform as well as the native species in the habitat of Oe. conioides. We hypothesize that selection against immigrants is the most important component of reproductive isolation between the two species, and that selection against immigrants is the result of exposure to cold and wind in the tidal habitat of Oe. conioides and to herbivory in the habitat of Oe. aquatica. These results are congruent with a role for differential ecological selection in the formation and maintenance of these two species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 526–535.     

山西桑干河流域湿地植被优势种群分布格局研究

根据野外调查的样地资料,应用方差/均值比（扩散系数）的t检验、聚集指数、平均拥挤度、聚块性指数、Green指数和负二项分布的聚集强度等方法,研究了山西桑干河流域湿地植被优势种群的分布格局并分析格局形成的原因,并用相关分析比较了6个指数间的关系,结果表明：这些植物的分布格局皆属于聚集分布,但它们的聚集强度有差异。聚集强度最大的是沙棘和柽柳,聚集强度最小的是小车前和车前,由于它们均分布于河漫滩或者河床地势较高处,土壤水分含量较低,仅仅在洪水季节短时间有积水存在,不是典型的湿地植物。典型的水生植物泽泻、莎草、轮藻、芦苇、香蒲、慈姑,湿生植物水葱、藨草等,它们的聚集强度居中,对水分有着较强的依赖性,均分布于流水较缓的河道,或者濒临河道的积水处。此外,多年生的莎草、芦苇、香蒲、水葱、藨草等主要依靠无性繁殖扩大种群分布范围和密度,这也是它们服从聚集分布的原因之一。本研究还说明方差/均值比率法是一种较好的分析种群分布格局的方法。     

Population genetic structure of native versus naturalized sympatric shrub willows (Salix; Salicaceae)

Vegetative propagation of an introduced species can contribute significantly to its ability to spread and become naturalized, potentially in competition with native species. This study focused on the naturalization of a willow shrub, Salix purpurea, which was introduced to the United States from Europe and is commonly sympatric with the native shrub willow, S. eriocephala. Both species are capable of vegetative and sexual reproduction, but little is known about their relative frequency, nor the impact of clonal propagation on population-level genetic diversity. We analyzed genotypes at several microsatellite loci in 993 individuals belonging to 30 subpopulations of S. eriocephala and 28 subpopulations of S. purpurea in areas of sympatry across three watersheds to compare their genetic diversity and genetic structure. Our results revealed six subpopulations of S. purpurea containing plants with identical multilocus genotypes, while clonal individuals were rare among S. eriocephala populations. These species are dioecious with relatively high levels of heterozygosity, but S. eriocephala had much higher allelic diversity and genotypic diversity than did S. purpurea. These results strongly suggest that vegetative propagation has contributed to the naturalization of S. purpurea and has resulted in higher levels of genetic differentiation among S. purpurea populations than among native S. eriocephala populations.     

Rapid growth of a Eurasian haplotype of <Emphasis Type="Italic">Phragmites australis</Emphasis> in a restored brackish marsh in Louisiana,USA

While numerous studies have documented patterns of invasion by non-indigenous plant species, few have considered the invasive properties of non-native genotypes of native species. Characteristics associated with specific genotypes, such as tolerance to disturbance, may mistakenly be applied to an entire species in the absence of genetic information, which consequently may affect management decisions. We report here on the incidence and growth of an introduced lineage of Phragmites australis in the Gulf of Mexico coastal zone of Louisiana. P. australis was collected from nine separate locations for inclusion in a series of growth experiments. Chloroplast DNA analysis indicated that specimens collected from four locations in the Mississippi River Delta represented the introduced Eurasian haplotype; the remainder represented the gulf coast haplotype. Three distinct genotypes, or clones, were identified within each haplotype via analysis using amplified fragment length polymorphisms, which also revealed reduced genetic diversity of the gulf coast clones compared to the Eurasian clones. Clones of each haplotype were planted along with three other native macrophytes at similar densities in a restored brackish marsh and monitored for growth. After 14 months, the Eurasian haplotype had spread vegetatively to cover about 82% of the experimental plots, more than four times the coverage (18%) of the gulf coast haplotype. Thus, the use of P. australis plantings for wetland restoration should consider the genetic lineage of plants used since our results indicate the potential of the Eurasian haplotype to grow rapidly at newly restored sites. This rapid growth may limit the establishment of more slowly growing native species.     

Intimately linked or hardly speaking? The relationship between genotype and environmental gradients in a Louisiana Iris hybrid population

Several models of hybrid zone evolution predict the same spatial patterns of genotypic distribution whether or not structuring is due to environment-dependent or -independent selection. In this study, we tested for evidence of environment-dependent selection in an Iris fulva x Iris brevicaulis hybrid population by examining the distribution of genotypes in relation to environmental gradients. We selected 201 Louisiana Iris plants from within a known hybrid population (80 m x 80 m) and placed them in four different genotypic classes (I. fulva, I. fulva-like hybrid, I. brevicaulis-like hybrid and I. brevicaulis) based on seven species-specific random amplified polymorphic DNA (RAPD) markers and two chloroplast DNA haplotypes. Environmental variables were then measured. These variables included percentage cover by tree canopy, elevation from the high water mark, soil pH and percentage soil organic matter. Each variable was sampled for all 201 plants. Canonical discriminant analysis (CDA) was used to infer the environmental factors most strongly associated with the different genotypic groups. Slight differences in elevation (-0.5 m to +0.4 m) were important for distinguishing habitat distributions described by CDA, even though there were no statistical differences between mean elevations alone. I. brevicaulis occurred in a broad range of habitats, while I. fulva had a narrower distribution. Of all the possible combinations, I. fulva-like hybrids and I. brevicaulis-like hybrids occurred in the most distinct habitat types relative to one another. Each hybrid class was not significantly different from its closest parent with regard to habitat occupied, but was statistically unique from its more distant parental species. Within the hybrid genotypes, most, but not all, RAPD loci were individually correlated with environmental variables. This study suggests that, at a very fine spatial scale, environment-dependent selection contributed to the genetic structuring of this hybrid zone.     

Sexual reproduction of cordgrass hybrids (Spartina foliosa x alterniflora) invading tidal marshes in San Francisco Bay

We genetically analysed cordgrass plants and seedlings throughout the San Francisco, California, USA, estuary and found that hybrids between exotic Spartina alterniflora and native Spartina foliosa are the principal cordgrass invaders and colonizers. We hypothesized that this was due to higher seed set and siring ability by hybrids relative to the native species; too few alien parents remained in San Francisco Bay for our comparative studies. Hybrid seed comprised 91% to 98% of that set in the marsh study plants over the 2 years of the study. Total viable pollen production by hybrid plants was 400 times that of the native plants. Seed and pollen production were highly skewed towards a few hybrid genotypes. In addition to seed produced by hybrid plants, hybrid seed was produced by S. foliosa due to hybrid backcrossing. While the greatest advantage for hybrids was in pollen and seed production, hybrid seeds germinated, and seedlings survived and grew as well or better than the native species. As native S. foliosa becomes increasingly rare, hybrid seed floating on the tides will predominate, overwhelming recruitment sites and resulting in further colonization by hybrids. In an evolutionary context, hybrids with exceptional pollen and seed production will be initially favoured by natural selection, leading to the evolution of even more fertile hybrid genotypes.     

Genetic Diversity in Wild Grapes Native to China Based on Randomly Amplified Polymorphic DNA (RAPD) Analysis

The use of the RAPD technique was investigated on a set of 73 genotypes of 18 wild grape species native to China, and one interspecific hybrid, seven Vitis vinifera L. cultivars, one rootstock cultivar and one strain of V. riparia L. Genetic diversity among these grapes was investigated based on RAPD analysis. The screening of 280 decamer oligonucleotides allowed the selection of 20 primers used for the analysis. A total of 191 RAPD markers were produced from the 20 selected primers. Relationships among the 83 clones or accessions based on their genetic distances were clustered using unweighted pair-group method arithmetic average (UPGMA) analysis in a dendrogram. Twenty-two clusters which fortunately adapted to 22 grape species level were clearly resolved on the dendrogram. The 18 wild grape species native to China were grouped into ten subclusters. The largest distance was found between V. riparia L., V. vinifera L., interspecific hybrid ( V. vinifera L.× V. larbrusca L.) and the wild grapes native to China. Among the wild grapes native to China, the largest distance was found between V. hancockii Hance and the other wild species. V. qinlingensis P.C.He was the second. Large genetic variation occurred among the different flower-type clones in one species.     

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.     

Genetic variation in photosynthetic characteristics among invasive and native populations of reed canarygrass (Phalaris arundinacea)

With the extensive spread of invasive species throughout North America and Europe there is an urgent need to better understand the morphological and physiological characteristics of successful invasive plants and the evolutionary mechanisms that allow introduced species to become invasive. Most ecological studies have focused on morphological differences and changes in community dynamics, and physiological studies have typically explored the differences between native and invasive species. In this study, 15 different genotypes of Phalaris arundinacea from both its native (European) and invasive (North American) range were grown in a common garden experiment to monitor the physiological differences between native and invasive genotypes. Here we present data that suggests high variability exists in the physiological traits among genotypes of P. arundinacea, yet genotypes from the native range are not necessarily physiologically inferior to the hybridized invasive genotypes. Previous work has shown that multiple introductions of P. arundinacea from various European locations to the United States resulted in numerous hybridization events, yielding more genetic variability and phenotypic plasticity in the invasive range. Of the genotypes studied, both morphological and physiological traits of genotypes with French origin were significantly different from the plants from the Czech Republic, North Carolina, and Vermont. The lack of clear differences between native and invasive genotypes indicates that physiological traits may be highly conserved in P. arundinacea and enhanced photosynthetic rates are not indicative of successful invasive genotypes. Instead, morphological traits and defensive secondary compound metabolism may play a more important role in the success of P. arundinacea within its invasive range, and patterns of genetic variation in physiological traits between invasive and native range may be more important than the mean traits of each region when explaining reed canarygrass’ invasive potential in North America.     

GENETIC STRUCTURE OF COEXISTING SEXUAL AND CLONAL SUBPOPULATIONS IN A FRESHWATER SNAIL (POTAMOPYRGUS ANTIPODARUM)

We examined clonal diversity and the distribution of both clonal and sexual genotypes in a single population of freshwater snails (Potamopyrgus antipodarum) in which diploid sexual individuals and triploid parthenogens coexist. A genetic analysis of individuals from three habitat zones in Lake Alexandrina, New Zealand revealed extremely high clonal diversity: 165 genotypes among 605 clonal individuals. The frequency of triploid clonal individuals increased with increasing depth in the lake, and most of the individual clones were habitat specific, suggesting that differences among habitats are important in structuring the clonal subpopulation. There were also high levels of clonal diversity within habitats, suggesting frequent origins of habitat-specific clones. In contrast, diploid sexual individuals were proportionately more common in the shallow regions of the lake (where infection by trematode larvae is highest), and there was no significant spatial structure in the sexual subpopulation. We suggest that habitat specialization by clones, as well as parasite-mediated selection against common clones, are important factors affecting the structure of this mixed population of sexual and clonal snails.     

Isolation and characterization of microsatellite markers in the rare clonal plant, Spiraea virginiana (Rosaceae)

? Premise of the study: Microsatellite markers were developed in Spiraea virginiana, a federally threatened native shrub found along stream banks, to identify clonal genotypes and measure population genetic variability. ? Methods and Results: Eleven primer sets were developed using a non-radioactive protocol. These revealed a moderate level of genetic variation, as indicated by the number of alleles per locus (range = 1-4) and an average observed heterozygosity of 0.595. Select loci also amplified successfully in the related species Spiraea japonica. ? Conclusion: Development of the markers described here is critical for the genetic identification of clonal plants as a first step in demographic analyses, and is necessary for the future conservation of this rare species. Amplification of the markers in S. japonica suggests their potential utility in research regarding this species.     

Spatial genetic structure and clonal diversity in an alpine population of Salix herbacea (Salicaceae)

BACKGROUND AND AIMS: Many alpine plant species combine clonal and sexual reproduction to minimize the risks of flowering and seed production in high mountain regions. The spatial genetic structure and diversity of these alpine species is strongly affected by different clonal strategies (phalanx or guerrilla) and the proportion of generative and vegetative reproduction. METHODS: The clonal structure of the alpine plant species Salix herbacea was investigated in a 3 x 3 m plot of an alpine meadow using microsatellite (simple sequence repeat; SSR) analysis. The data obtained were compared with the results of a random amplified polymorphic DNA (RAPD) analysis. KEY RESULTS: SSR analysis, based on three loci and 16 alleles, revealed 24 different genotypes and a proportion of distinguishable genotypes of 0.18. Six SSR clones were found consisting of at least five samples, 17 clones consisting of more than two samples and seven single genotypes. Mean clone size comprising at least five samples was 0.96 m(2), and spatial autocorrelation analysis showed strong similarity of samples up to 130 cm. RAPD analysis revealed a higher level of clonal diversity but a comparable number of larger clones and a similar spatial structure. CONCLUSIONS: The spatial genetic structure as well as the occurrence of single genotypes revealed in this study suggests both clonal and sexual propagation and repeated seedling recruitment in established populations of S. herbacea and is thus suggestive of a relaxed phalanx strategy.     

Herbivory and local genetic differentiation in natural populations of Arabidopsis thaliana (Brassicaceae)

To explore genetic variation in defence against the natural herbivores of Arabidopsis thaliana, we transplanted genotypes between a dune habitat and inland habitat in both of which A. thaliana occurred naturally. In previous years we had observed that the specialist weevils Ceutorhynchus atomus and C. contractus (Curculionidae) fed conspicuously on flowers and fruits of A. thaliana in the dunes, while these weevils were always rare in inland habitats. Taking all plants together, total fruit damage was indeed much higher in our experimental plots in the dune habitat (59.7%) relative to the inland garden habitat (18.9%). Within a habitat, additional differences existed between plants of different origins, pointing to genetic differences in ecologically relevant characters; plants of inland origin flowered a week earlier, grew better and produced more fruits than plants of dune origin. However, plants of inland origin experienced more total fruit damage by the specialist weevils (75.4%) than plants of dune origin (44.0%) when the two types grew side by side in the dune habitat. Escape from herbivory gives dune genotypes an advantage in their native habitat, whereas stronger growth and higher survival gives inland genotypes an edge under garden conditions.     

Adaptation of Senecio vulgaris (Asteraceae) to ruderal and agricultural habitats

Adaptation of the annual plant Senecio vulgaris to ruderal and agricultural habitats was investigated. We expected S. vulgaris to be adapted to the agricultural habitat through nutrient-specific differentiation of relatively few genotypes responding to the generally high homogenous nutrient levels at the agricultural habitat caused by constant fertilization. To assess adaptation of S. vulgaris, vegetative and reproductive responses of seed families from various populations of agricultural and ruderal habitats, grown in the greenhouse at high and low nutrient levels, were compared. Data were analyzed with a three-level nested ANOVA based on the levels habitat, population, and family. A significant habitat effect indicated that S. vulgaris from ruderal and agricultural habitats were genetically different with plants from the agricultural habitat having larger leaves and a higher reproduction. A significant habitat by nutrient effect showed a stronger response of reproduction to nutrients at the agricultural habitat, suggesting that genetic differentiation among habitats is nutrient-specific. Contrary to expectations, only the agricultural habitat showed genetic diversity of S. vulgaris. Results suggest that nutrient levels at the agricultural habitat are more heterogeneous as generally proposed leading to a relatively high genetic variation.     

Genetic variation of introduced Hawaiian and native Costa Rican populations of an invasive tropical shrub, Clidemia hirta (Melastomataceae)

Clidemia hirta is one of the most common woody invasive plants in mesic to wet forests in Hawaii, where it was introduced around 1940. The species is relatively uncommon by comparison in its native range of Central and South America and some Caribbean Islands. We examined genetic variation in allozymes of 20 C. hirta populations on four Hawaiian Islands to determine the introduction history. For comparison, we measured genetic variation in 20 native populations across Costa Rica. Mean levels of genetic variation in Hawaiian and Costa Rican populations were low compared to other woody or introduced plants (11.5-12.5% polymorphic loci, 2.05-2.50 alleles per polymorphic locus, and 0.045-0.063 expected heterozygosity). Most genetic diversity was held within rather than among populations in both areas (G(ST) = 0.120 and 0.271 in Hawaii and Costa Rica, respectively). Hawaiian populations had a high degree of genetic similarity, and no genetic differentiation was found among the four Hawaiian Islands sampled. These patterns of genetic variation in Hawaii suggest that no intraspecific hybridization of genotypes from different parts of the native range has occurred and that introductions to the different islands came from the same or similar source populations. The low levels of genetic diversity in parts of both the native and introduced ranges suggest that genetic variation is unrelated to invasiveness in C. hirta.     

Microgeographic genetic structure and gene flow in Hibiscus moscheutos (Malvaceae) populations

Microgeographic genetic variation in populations of a wetland macrophyte, Hibiscus moscheutos L. (Malvaceae), was investigated using allozyme polymorphism. The species is a self-compatible insect-pollinated perennial, and seeds are water dispersed (hydrochory). Six hundred plants were analyzed from eight brackish and two freshwater populations within the Rhode River watershed/estuarine system. The genetic structure of the populations was assessed by fixation indices and spatial autocorrelation analyses. The degree of genetic differentiation among sites and gene flow between all paired combinations of sites (M ) was analyzed using three hypothetical gene flow models. Fixation indices indicated almost complete panmixia within populations, and spatial autocorrelations showed that genotypes were randomly distributed within sites, most likely the result of water dispersal of seeds. Allele frequencies were significantly different among sites, and estimated FST indicated moderate genetic differentiation (_ = 0.062). Genetic differences between populations were mostly explained by a gene flow model that accounted for the location of populations relative to the tidal stream. The importance of hydrochory in affecting spatial genetic structure was thus suggested both within and among H. moscheutos populations.     

Hybridization between introduced smooth cordgrass (Spartina alterniflora; Poaceae) and native California cordgrass (S. foliosa) in San Francisco Bay, California, USA

Introduced Spartina alterniflora (smooth cordgrass) is rapidly invading intertidal mudflats in San Francisco Bay, California. At several sites, S. alterniflora co-occurs with native S. foliosa (California cordgrass), a species endemic to California salt marshes. In this study, random amplified polymorphic DNA markers (RAPDs) specific to each Spartina species were identified and used to test for hybridization between the native and introduced Spartina species in the greenhouse and in the field. Greenhouse crosses were made using S. alterniflora as the pollen donor and S. foliosa as the maternal plant, and these crosses produced viable seeds. The hybrid status of the crossed offspring was confirmed with the RAPD markers. Hybrids had low self-fertility but high fertility when back-crossed with S. foliosa pollen. Hybrids were also found established at two field sites in San Francisco Bay; these hybrids appeared vigorous and morphologically intermediate between the parental species. Field observations suggested that hybrids were recruiting more rapidly than the native S. foliosa. Previous work identified competition from introduced S. alterniflora as a threat to native S. foliosa. In this study, we identify introgression and the spread of hybrids as an additional, perhaps even more serious threat to conservation of S. foliosa in San Francisco Bay.