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.     

Inbreeding depression in smooth cordgrass (Spartina alterniflora, Poaceae) invading San Francisco Bay

The magnitude of inbreeding depression in invading plant populations is often presumed to be small and of little consequence. The purpose of this study was to assess the magnitude of inbreeding depression in a pollen-limited, partially self-incompatible, invading plant population. The magnitude and timing of inbreeding depression were compared among ten maternal plants sampled from a population of smooth cordgrass (Spartina alterniflora) invading San Francisco Bay. Selfed and outcrossed progeny were compared for embryo abortion, survival of seedlings, and growth/survival at the end of the first growing season in three greenhouse environments. Estimates of inbreeding depression varied among environments, with competitive environment > high-nutrient environment > low-nutrient environment. Population-level estimates of inbreeding depression ranged from 0.61 to 0.81; however, maternal plants varied significantly in their magnitude of inbreeding depression, ranging from 0.1 to 0.97. The 95% confidence interval for inbreeding depression for some maternal plants included zero. There was a significant negative correlation between the overall magnitude of inbreeding depression and self-fertility rate among maternal plants. The few maternal plants with high self-fertility carried relatively little genetic load, and their selfed progeny are likely to survive on open mudflats. The noncompetitive, pollen-limited growing conditions associated with invasion may allow self-fertility to spread in this population.     

Characterization of microsatellite loci in Spartina species (Poaceae)

The cordgrasses in the genus Spartina have become model organisms for studying biological invasions from both ecological and genetic perspectives. Here we characterize 11 disomic loci in Spartina alterniflora that show promise for population studies and for studying hybridization events between S. alterniflora and S. foliosa. Comparisons among invasive and native S. alterniflora populations showed that levels of allelic variation are lower in invasive populations. In addition, nearly all loci that amplified in S. foliosa populations and in a swarm of S. alterniflora×foliosa hybrids were polymorphic. We also found that several loci amplified successfully in other Spartina species.     

Effects of drainage and soil organic content on growth of Spartina alterniflora (Poaceae) in an artificial salt marsh mesocosm

An artificial salt marsh mesocosm was constructed using 680-L polypropylene tanks to determine the effect of soil drainage depth and organic content on growth and rhizome proliferation of the salt marsh smooth cordgrass Spartina alterniflora. Soil drainage depth had no effect on accumulation of aerial or subsurface plant tissue, but tanks that had 2.5% soil organic content supported enhanced aerial tissue and rhizome growth compared to tanks that contained sand alone. We propose a mathematical model for predicting the mass of photosynthetically significant leaf tissue without cutting and drying leaves. Implications of these findings for salt marsh creation projects are discussed.     

Extent and degree of hybridization between exotic (Spartina alterniflora) and native (S. foliosa) cordgrass (Poaceae) in California, USA determined by random amplified polymorphic DNA (RAPDs)

Spartina alterniflora, smooth cordgrass, native to the eastern USA, was introduced into south San Francisco Bay ≈ 25 years ago. It has spread by purposeful introduction of rooted plants and dispersal of seeds on the tides. Previous work suggested that S. alterniflora was competitively superior to the native California cordgrass, S. foliosa, and that the two species hybridized. The present study determined the spread of S. alterniflora and S. foliosa × alterniflora hybrids in California and examined the degree of hybridization. We used nuclear DNA markers diagnostic for each species to detect the parental species and nine categories of hybrids. The California coast outside San Francisco Bay contained only the native species. All hybrid categories exist in the Bay, implying that several generations of crossing have occurred and that hybridization is bidirectional. Hybrids were found principally near sites of deliberate introduction of the exotic species. Where S. alterniflora was deliberately planted, we found approximately equal numbers of S. alterniflora and hybrid individuals; S. foliosa was virtually absent. Marshes colonized by water-dispersed seed contained the full gamut of phenotypes with intermediate-type hybrids predominating. The proliferation of hybrids could result in local extinction of S. foliosa. What is more, S. alterniflora has the ability to greatly modify the estuary ecosystem to the detriment of other native species and human uses of the Bay. To the extent that they share these engineering abilities, the proliferation of cordgrass hybrids could grossly alter the character of the San Francisco Bay.     

互花米草的两面性及其生态控制

互花米草是美国大西洋沿岸的本土植物,其高生产力、高繁殖率、高抗性和致密发达的地下部分等生物学特性使它能在沿海地区的抗风防浪、保滩护岸、促淤造陆、修复湿地和固定二氧化碳等方面发挥很好的正生态效应;然而,正是这与生俱来的生物学优势使该物种在引种地区侵占光滩,很快形成茂盛的单种优势群落,体现出很强的入侵性,从而也具有相当大的负生态效应。本文综述了互花米草的正负生态效应,分别从以下4个方面展开:抗风防浪的功效与改变景观的负作用,促淤造陆的功效与改变河口水文格局的负作用,固碳效应与释放温室气体的负作用,为一些物种提供营养与栖息地的同时对其他物种栖息环境的破坏性。研究证实,互花米草的本土化倾向已比较突出,如在超过100年引种历史的美国西部维拉帕湾、引种历史近40年的旧金山湾,以及引种历史近30年的我国大丰滩涂等。对于互花米草的生态控制,应结合各地的自然、社会、经济状况,因地制宜,如在一些生态脆弱(开发过度、缺乏大型植被、蚀退现象严重)的沿海地区,防灾减灾任务艰巨,根除互花米草须谨慎;在充分利用互花米草抗风防浪和促淤造陆后,可以适时围垦,以达到某些区域大面积减除的目标;此外,综合治理与利用相结合是较为科学的举措,既充分利用互花米草生物质(加工生产新食品、保健品、生物气等),又有效控制其种子飘落和扩繁,是一项生态工程,也是一种具有生态经济效益的利国利民之举。     

互花米草黄酮含量分析及其生态学意义

互花米草原产于北美洲大西洋沿岸和墨西哥海湾,经引种和其他原因,已扩散到包括中国在内的多个国家和地区,并对被入侵地生态系统的结构和功能均造成了显著影响。有关该物种的入侵机制和控制策略是当前入侵生态学研究领域的热点。互花米草生物质材料中富含黄酮类物质,与该物种向海扩张的能力可能存在一定联系。鉴于黄酮广泛的应用价值,能否借助黄酮成分的开发带动生物质的收割,以达到有效控制该物种的目的,也是当前研究者所关注的问题。以芦丁为标准参照物,NaNO2-Al（ NO3）3法比较温室互花米草不同器官中总黄酮含量的差异,分析盐城滩涂互花米草越冬芽黄酮含量与所在地理位置之间的关系,采用L9（34）正交试验对干储互花米草叶总黄酮的提取工艺进行优化。互花米草叶片中黄酮类物质远高于根、根状茎、茎、叶鞘、颖稃及种子。盐城滩涂上互花米草越冬芽的黄酮含量随着由陆向海方向基本呈升高趋势。互花米草总黄酮最佳提取工艺为料液比1∶30,乙醇浓度70％,水浴温度80℃,水浴时间2h。互花米草植物材料中,尤其在叶部,黄酮含量十分丰富。该黄酮物质与互花米草在海岸带盐沼生态系统中的适应和向海扩张也存在正相关关系,可作为探索互花米草入侵机制的一个方向。利用本研究中所得优化提取工艺可从互花米草干储生物质中获得稳定的黄酮来源。配合黄酮经济成分的开发而推动生物质材料的收割利用,将成为互花米草生态控制的一项潜在策略。     

Evolution of a new ecotype of Spartina alterniflora (Poaceae) in San Francisco Bay, California, USA

We report the discovery and spread of a dwarf ecotype of Spartina alterniflora in San Francisco Bay. Relative to typical S. alterniflora, this dwarf ecotype has one-fifth the tiller height (～21 cm), tenfold the tiller density (～4000 tillers/m(2)), and is restricted to growth in the upper intertidal zone. Chromosome counts of the dwarfs are identical to typical smooth cordgrass (2n = 62), and smooth cordgrass-specific random amplified DNA markers confirm the species identity of the dwarf. Field-collected clonal fragments of the dwarf grown for 2 yr under high-nutrient conditions maintained the dwarf syndrome, as did plants grown from the seed of a dwarf. The dwarf condition is not caused by endophytic fungi. The first dwarf smooth cordgrass patch was discovered in 1991, and by 1996 five separate dwarf patches had appeared within 200 m of the original. Since 1991, total area covered by the dwarf ecotype has increased sixfold to 140 m(2). The ecological range of the dwarf smooth cordgrass ecotype is similar to that of S. patens, a competitor on the Atlantic coast. We suggest that the absence of S. patens from most of San Francisco Bay has allowed the dwarf ecotype of smooth cordgrass to survive and spread.     

Genetic evidence for hybridization between the native Spartina maritima and the introduced Spartina alterniflora (Poaceae) in South-West France: Spartina × neyrautii re-examined

 Spartina alterniflora, a perennial grass native to the North American Atlantic coast, was introduced during the 19th century in western Europe (Southern England and western France) where it hybridized with the native Spartina maritima. In England, the sterile hybrid S. × townsendii gave rise by chromosome doubling to the highly fertile allopolyploid Spartina anglica, which has now invaded many salt marshes and estuaries in western Europe, and has been introduced in several continents. In South-West France, another sterile hybrid was discovered in 1892 in the Bidassoa Estuary, and named Spartina × neyrautii. According to their morphology, some authors suggested that S. × neyrautii and S. × townsendii result from reciprocal crosses. During the 20th century, the hybridization site was severely disturbed, and surviving of S. × neyrautii was questioned. In this paper, various Spartina populations are investigated in the Basque region (France and Spain), and compared to the hybrid taxa formed in England (S. × townsendii and S. anglica). The samples were analyzed using molecular fingerprinting (RAPD and ISSR) and Chloroplast DNA sequence (trnL-trnT spacer, trnL intron and trnL-trnF spacer). In the Bidassoa estuary, a hybrid isolated clone has been found, that displays additive species-specific nuclear markers of S. maritima and S. alterniflora, and that is subsequently considered as a surviving clone of S. × neyrautii. The molecular analyses indicate that S. × neyrautii and S. × townsendii share the same maternal (S. alterniflora), and paternal (S. maritima) parental species, but also that the two independent hybridization events have involved different parental (nuclear) genotypes in England and in South-West France. Received July 12, 2002; accepted October 4, 2002 Published online: March 20, 2003     

Effect of Salt Stress on Seed Germination and EmbryoGrowth of Spartina alterniflora (Gramineae)

In this paper , the effect of NaCl concentration ( 1× 104 - 6× 104 mg/L) on the seed germination of smooth cordgrass ( Spartina alterniflora) was investigated using 1/2 Hoagland as the basic culture solution. The results indicated that the coleoptile and embryonal axis grew before the radicle and plumule in the stage of germination . Salinity level had no influence on the seed germination percentage when the salt concentration didn′t exceed 3×104 mg/L. The optimal salinity level for seed germination and embryo growth was 1×104 mg/L . The inhibit effect of salt concentration on the growth of embryo′s different part was different, and the length of radicle and plumule decreased significantly with the increase of salt concentration. However , the lesser inhibition on the embryonal axis and coleoptile was observed under the higher salt concentration, and thus the seedlings out of soil were ensured and the species was quickly adapted to the environment of inter- tidal zone .     

A comparison of the seed dormancy characteristics of Spartina patens and Spartina alterniflora (Poaceae)

Seed dormancy characteristics of Spartina alterniflora were delineated previously by Plyler and Carrick (American Journal of Botany, vol. 80, pp. 752–756, 1993). This study was undertaken in order to determine whether or not the dormancy characteristics of S. patens are similar. As in the previous study, the site of a dormancy mechanism was determined by assessing the germinability of surgically altered dormant seeds. Likewise, the effects of three growth-regulating substances (abscisic acid, fusicoccin, and gibberellic acid), prechilling, and light were studied by assessing the germinability of appropriately treated dormant seeds. Surgical injury to the endosperm, and to a lesser extent the scutellum, produced significant germination in S. patens, whereas only injury to the scutellum produced germination in S. alterniflora. Exogenously applied abscisic acid was highly effective in maintaining dormancy in injured seeds of both species. Gibberellic acid and fusicoccin were ineffective in promoting germination in S. patens, but fusicoccin was highly effective in breaking dormancy in 5. alterniflora. Prechilling was effective in breaking dormancy in S. patens but not in S. alterniflora. In S. patens, treatments that broke dormancy were only successful when seeds were exposed to light during the germination period. It is concluded that the dissimilarities in the dormancy mechanisms may be manifestations of the different environmental adaptations these closely related species display.     

Effect of oxygen availability and salinity on EARLY LIFE HISTORY STAGES OF SALT MARSH PLANTS. I. Different germination strategies of Spartina alterniflora and Phragmites australis (Poaceae)

Gradients in oxygen availability and salinity are among the most important environmental parameters influencing zonation in salt marsh communities. The combined effects of oxygen and salinity on the germination of two salt marsh grasses, Spartina alterniflora and Phragmites australis, were studied in growth chamber experiments. Germination of both species was initiated by emergence of the shoot and completed by root emergence. Percentage S. alterniflora germination was reduced at high salinity (40 g NaCl/L) and in decreased oxygen (5 and 2.5%). In 0% oxygen shoots emerged, but roots did not. P. australis germination was reduced at a lower salinity (25 g NaCl/L) than S. alterniflora, and inhibited at 40 g NaCl/L and in anoxia. However, a combination of hypoxia (10 and 5% O2) and moderate salinity (5 and 10 g NaCl/L) increased P. australis germination. When bare areas in the salt marsh are colonized, the different germination responses of these two species to combinations of oxygen and salt concentrations are important in establishing their initial zonation. In high salinity wetlands S. alterniflora populates the lower marsh and P. australis occupies the high marsh at the upland boundary.     

互花米草与海三棱藨草的生长特征和相对竞争能力

外来物种互花米草(Spartinaalterniflora)已对我国特有种海三棱藨草 (Scirpusmariqueter)发生了竞争取代。作者通过在长江口崇明东滩湿地取样和盆栽竞争实验,比较了二者的主要生长特征和种间相对竞争能力。结果表明:无论是先锋种群还是成熟种群,互花米草的高度、盖度、地上生物量、地下生物量和平均每花序种子数都显著大于海三棱藨草;海三棱藨草的密度和单位面积结实枝条数显著大于互花米草。两种植物成熟种群的单位面积种子产量和种子的萌发率没有显著差异。种内竞争和种间竞争显著降低了两种植物的平均每株产生的无性小株数、结实株数、地上生物量和地下球茎数(海三棱藨草产生球茎),互花米草的种间竞争能力 (相对邻里效应指数 )显著大于海三棱藨草。互花米草在生长上的优势可能导致其在种间竞争上的优势,从而使互花米草能够在海三棱藨草群落中成功入侵。     

长江口互花米草生长区大型底栖动物的群落特征

2005年对长江口潮滩湿地互花米草(Spartina alterniflora)生长区不同季节大型底栖动物群落特征的研究表明:长江口互花米草生长区的大型底栖动物有21种,其中甲壳纲10种、多毛纲5种、腹足纲5种、辨鳃纲1种.主要种类有拟沼螺(Assiminea sp.)、背蚓虫(Notomastus latericeus)、尖锥拟蟹守螺(Cerithidea largillierli)、中国绿螂(Glauconome chinensis)、钩虾(Gammaridae sp.)等.食性功能群均以碎屑食者和植食者为主.大型底栖动物平均栖息密度为(650.5±719.2)个/m2,标准误主要是由于北湖的拟沼螺密度很大.栖息密度从大到小的顺序为沿河口梯度从内到外分布的北湖边滩、崇明东滩、金山卫边滩.大型底栖动物群落分布不均匀,沿河口梯度变化存在明显的空间差异.栖息密度和物种多样性在夏季最高,冬季最低.大型底栖动物平均生物量为(20.8±6.1)g/m2,季相变化为夏季＞秋季＞春季＞冬季.BIO-ENV分析表明沉积物粒径和盐沼高度是大型底栖动物群落特征的主要影响因素.不同研究结果的差异除了时空因素外可能与互花米草的种群动态有关.加强不同时间尺度的研究有助于正确评价互花米草对大型底栖动物的影响.     

闽江河口互花米草不同入侵年限湿地土壤磷赋存形态

选择闽江河口鳝鱼滩互花米草不同入侵年限湿地(SA₁,5～6年;SA₂,8～10年;SA₃,12～14年)为对象,基于时空互代法研究了互花米草入侵对湿地土壤磷赋存形态的影响。结果表明: 不同入侵年限互花米草湿地土壤中难分解态磷(HCl-Pi、Residual-P)含量的占比最高(46.4%～46.7%),中等易分解态磷(NaOH-Pi、NaOH-Po、Sonic-Pi)次之(40.0%～44.0%),而易分解态磷(Resin-Pi、NaHCO₃-Pi、NaHCO₃-Po)最低(9.5%～13.3%)。随着互花米草入侵年限的增加,湿地土壤磷赋存形态及其空间分布特征也发生明显改变,中等易分解态磷、难分解态磷和总磷含量整体上均呈增加趋势,而易分解态磷含量呈降低趋势。相对于SA₁,SA₂的中等易分解态磷、难分解态磷和总磷含量分别增加了11.5%、9.7%和10.5%,而SA₃分别增加了24.8%、13.2%和13.5%。互花米草入侵年限的增加显著改变了湿地土壤磷的赋存形态特征,这主要取决于电导率(EC)、pH值及粒度等关键因子的变化。近年来,闽江河口开展的互花米草定期刈割治理在一定程度上减少了残体向土壤归还磷养分的能力,降低了土壤中易分解态磷的养分供应。     

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.     

外来种互花米草对盐沼土壤微生物多样性的影响——以江苏滨海为例

研究了江苏滨海外来种互花米草的生长对潮间带土壤微生物特征的影响,结果显示:与原有光滩相比,互花米草的大面积生长,使当地潮间带土壤微生物量增加,并随植被的生长状况发生变化,潮间带土壤微生物对碳和营养物质的利用相对较少;同时微生物生理功能群中占优势的活动组分发生了变化,组成可能更复杂。土壤微生物群落生理功能多样性(CLPP)分析结果显示,潮间带土壤微生物群落对碳源底物的利用能力不高,所利用碳源以糖类、氨基酸类、羧酸类占优势;其中互花米草盐沼微生物利用的碳源种类多于光滩,并且不同季节土壤微生物群落碳源利用类型存在差异;潮间带土壤微生物在不同的季节中其生理功能群组成发生着变化,互花米草盐沼的土壤微生物在植被生长减弱时具有与光滩相近的代谢类群,在生理功能上具有一定的相似性;当植被生长旺盛时,其代谢类群增多,代谢强度升高,微生物群落功能多样性增加。外来种互花米草在滨海潮间带的大面积生长,改善了土壤理化性质,为该处土壤微生物提供了不同的碳源,增强了土壤微生物的活动,改变了土壤微生物群落生理功能结构。     

Variation in self-fertility and the reproductive advantage of self-fertility for an invading plant (Spartina alterniflora)

The factors responsible for the reproductive success or failure of individuals in small, founding populations have received little attention. Previous work on a small population of smooth cordgrass (Spartina alterniflora) invading San Francisco Bay, California found that most clones flower prolifically but set little or no seed, while a few clones have high rates of viable seed set, producing most of the seeds in the population. This study first identifies recruitment from seeds as the main source of new smooth cordgrass plants during invasion and then tests the influence of growing conditions and pollination treatment on viable seed set among clones established in San Francisco Bay. Field transplants indicated that a clone's seed set rate was not strongly dependent on its site of establishment. Low and high nutrient greenhouse treatments also had little effect on viable seed set rates within most clones. In contrast, pollination treatment (self-pollination or outcrossing) had a major effect on viable seed set rates. Most clones had high seed set rates after outcross-pollination, but clones varied widely in their selfing capacity. Zero or low viable self-seed set rates were most common; however, a few clones had high viable self-seed set rates, comparable to outcross seed set rates. A clone's selfing capacity was significantly correlated across years (r=0.89, P<0.001), and capacity to set viable self-pollinated seeds in the greenhouse was significantly correlated with the clone's rate of viable seed set in the field . In this growing population where cross-pollination is limited, only the clones with high selfing ability had high viable seed set rates in the field. Among primarily outcrossing plant invaders, variation in self-fertility among individuals may be a common phenomenon, with important implications for genetic differentiation, effective population size and patterns of spatial spread during an invasion.     

A native dasyurid predator (common planigale,Planigale maculata) rapidly learns to avoid a toxic invader

Abstract Interactions between invasive species and native fauna afford a unique opportunity to examine interspecific encounters as they first occur, without the complications introduced by coevolution. In northern Australia, the continuing invasion of the highly toxic cane toad Bufo marinus poses a threat to many frog‐eating predators. Can predators learn to distinguish the novel toxic prey item from native prey (and thus, avoid being poisoned), or are longer‐term genetically based changes to attack behaviour needed before predators can coexist with toads? To predict the short‐term impact of cane toads on native predators, we need to know the proportion of individuals that will attack toads, the proportion surviving the encounter, and whether surviving predators learn to avoid toads. We quantified these traits in a dasyurid (common planigale, Planigale maculata) that inhabits tropical floodplains across northern Australia. Although 90% of naïve planigales attacked cane toads, 83% of these animals survived because they either rejected the toad unharmed, or killed and consumed the prey snout‐first (thereby avoiding the toxin‐laden parotoid glands). Most planigales showed one‐trial learning and subsequently refused to attack cane toads for long time periods (up to 28 days). Toad‐exposed planigales also avoided native frogs for up to 9 days, thereby providing an immediate benefit to native anurans. However, the predators gradually learnt to use chemical cues to discriminate between frogs and toads. Collectively, our results suggest that generalist predators can learn to distinguish and avoid novel toxic prey very rapidly – and hence, that small dasyurid predators can rapidly adapt to the cane toad invasion. Indeed, it may be feasible to teach especially vulnerable predators to avoid cane toads before the toads invade, by deploying low‐toxicity baits that stimulate taste‐aversion learning.