Strong genetic differentiation in the invasive annual grass <Emphasis Type="Italic">Bromus tectorum</Emphasis> across the Mojave–Great Basin ecological transition zone

Bromus tectorum, an inbreeding annual grass, is a dominant invader in sagebrush steppe habitat in North America. It is also common in warm and salt deserts, displaying a larger environmental tolerance than most native species. We tested the hypothesis that a suite of habitat-specific B. tectorum lineages dominates warm desert habitats. We sampled 30 B. tectorum Mojave Desert and desert fringe populations and genotyped 10–26 individuals per population using 69 single nucleotide polymorphic (SNP) markers. We compared these populations to 11 Great Basin steppe and salt desert populations. Populations from warm desert habitats were dominated by members of two haplogroups (87 % of individuals) that were distinct from haplogroups common in Great Basin habitats. We conducted common garden studies comparing adaptive traits and field performance among haplogroups typically found in different habitats. In contrast to the haplogroup abundant in sagebrush steppe, warm desert haplogroups generally lacked a vernalization requirement for flowering. The most widespread warm desert haplogroup (Warm Desert 1) also had larger seeds and a higher root:shoot ratio than other haplogroups. In the field, performance of warm desert haplogroups was dramatically lower than the sagebrush steppe haplogroup at one steppe site, but one warm desert haplogroup performed as well as the steppe haplogroup under drought conditions at the other site. Our results suggest that B. tectorum succeeds in widely disparate environments through ecotypic variation displayed by distinct lineages of plants. Accounting for this ecotypic variation is essential in modeling its future distribution in response to climate change.     

Exotic plants increase and native plants decrease with loss of foundation species in sagebrush steppe

Dominant plant species, or foundation species, are recognized to have a disproportionate control over resources in ecosystems, but few studies have evaluated their relationship to exotic invasions. Loss of foundation species could increase resource availability to the benefit of exotic plants, and could thereby facilitate invasion. The success of exotic plant invasions in sagebrush steppe was hypothesized to benefit from increased available soil water following removal of sagebrush (Artemisia tridentata), a foundation species. We examined the effects of sagebrush removal, with and without the extra soil water made available by exclusion of sagebrush, on abundance of exotic and native plants in the shrub steppe of southern Idaho, USA. We compared plant responses in three treatments: undisturbed sagebrush steppe; sagebrush removed; and sagebrush removed plus plots covered with “rainout” shelters that blocked winter-spring recharge of soil water. The third treatment allowed us to examine effects of sagebrush removal alone, without the associated increase in deep-soil water that is expected to accompany removal of sagebrush. Overall, exotic herbs (the grass Bromus tectorum and four forbs) were 3–4 times more abundant in shrub-removal and 2 times more abundant in shrub-removal + rainout-shelter treatments than in the control treatment, where sagebrush was undisturbed. Conversely, native forbs were only about half as abundant in shrub removal compared to control plots. These results indicate that removal of sagebrush facilitates invasion of exotic plants, and that increased soil water is one of the causes. Our findings suggest that sagebrush plays an important role in reducing invasions by exotic plants and maintaining native plant communities, in the cold desert we evaluated.     

Ecological genetics of vernalization response in Bromus tectorum L. (Poaceae)

BACKGROUND AND AIMS: Bromus tectorum (cheatgrass or downy brome) is an exotic annual grass that is dominant over large areas of former shrubland in western North America. To flower in time for seed production in early summer, B. tectorum plants generally require vernalization at winter temperatures, either as imbibed seeds or as established seedlings. METHODS: Variation in response to increasing periods of vernalization as seeds or seedlings for progeny of ten full-sib families from each of four B. tectorum populations from contrasting habitats was studied. KEY RESULTS: As vernalization was increased from 0 to 10 weeks, the proportion of plants flowering within 20 weeks increased, weeks to initiation of flowering decreased, and seed yield per plant increased, regardless of whether plants were vernalized as seeds or seedlings. Most of the variation was accounted for by differences among populations. Plants of the warm desert population flowered promptly even without vernalization, while those of the cold desert, foothill and montane populations showed incremental changes in response variables as a function of vernalization period. Populations differed in among-family variance, with the warm desert population generally showing the least variance and the cold desert population the most. Variation among populations and among families within populations decreased as vernalization period increased, whereas the non-genetic component of variance showed no such pattern. CONCLUSIONS: Variation in vernalization response was found to be adaptively significant and apparently represents the result of contrasting selection regimes on a range of founder genotypes.     

Conversion of sagebrush shrublands to exotic annual grasslands negatively impacts small mammal communities

Aim The exotic annual cheatgrass (Bromus tectorum) is fast replacing sagebrush (Artemisia tridentata) communities throughout the Great Basin Desert and nearby regions in the Western United States, impacting native plant communities and altering fire regimes, which contributes to the long‐term persistence of this weedy species. The effect of this conversion on native faunal communities remains largely unexamined. We assess the impact of conversion from native perennial to exotic annual plant communities on desert rodent communities. Location Wyoming big sagebrush shrublands and nearby sites previously converted to cheatgrass‐dominated annual grasslands in the Great Basin Desert, Utah, USA. Methods At two sites in Tooele County, Utah, USA, we investigated with Sherman live trapping whether intact sagebrush vegetation and nearby converted Bromus tectorum‐dominated vegetation differed in rodent abundance, diversity and community composition. Results Rodent abundance and species richness were considerably greater in sagebrush plots than in cheatgrass‐dominated plots. Nine species were captured in sagebrush plots; five of these were also trapped in cheatgrass plots, all at lower abundances than in the sagebrush. In contrast, cheatgrass‐dominated plots had no species that were not found in sagebrush. In addition, the site that had been converted to cheatgrass longer had lower abundances of rodents than the site more recently converted to cheatgrass‐dominated plots. Despite large differences in abundances and species richness, Simpson’s D diversity and Shannon‐Wiener diversity and Brillouin evenness indices did not differ between sagebrush and cheatgrass‐dominated plots. Main conclusions This survey of rodent communities in native sagebrush and in converted cheatgrass‐dominated vegetation suggests that the abundances and community composition of rodents may be shifting, potentially at the larger spatial scale of the entire Great Basin, where cheatgrass continues to invade and dominate more landscape at a rapid rate.     

Population genetic analysis of Bromus tectorum (Poaceae) indicates recent range expansion may be facilitated by specialist genotypes

? Premise of the study: The mechanisms for range expansion in invasive species depend on how genetic variation is structured in the introduced range. This study examined neutral genetic variation in the invasive annual grass Bromus tectorum in the Intermountain Western United States. Patterns of microsatellite (SSR) genotype distribution in this highly inbreeding species were used to make inferences about the roles of adaptively significant genetic variation, broadly adapted generalist genotypes, and facultative outcrossing in the recent range expansion of B. tectorum in this region. ? Methods: We sampled 20 individuals from each of 96 B. tectorum populations from historically and recently invaded habitats throughout the region and used four polymorphic SSR markers to characterize each individual. ? Key results: We detected 131 four-locus SSR genotypes; however, the 14 most common genotypes collectively accounted for 79.2% of the individuals. Common SSR genotypes were not randomly distributed among habitats. Instead, characteristic genotypes sorted into specific recently invaded habitats, including xeric warm and salt desert as well as mesic high-elevation habitats. Other SSR genotypes were common across a range of historically invaded habitats. We observed very few heterozygous individuals (0.58%). ? Conclusions: Broadly adapted, generalist genotypes appear to dominate historically invaded environments, while recently invaded salt and warm desert habitats are dominated by distinctive SSR genotypes that contain novel alleles. These specialist genotypes are not likely to have resulted from recombination; they probably represent more recent introductions from unknown source populations. We found little evidence that outcrossing plays a role in range expansion.     

Ecological genetics of Bromus tectorum

Summary By incorporating demographic analyses of fitness components (e.g., survival and reproduction) within a reciprocal sowing design, we tested for 3 consecutive years whether local adaptation has occurred in the alien grass Bromus tectorum (cheatgrass) within 7 habitats along an environmental gradient from arid steppe to subalpine forest in the Intermontain Region of western North America. Patterns of emergence and survival were strongly influenced by the local environment. In terms of survival, expression of significant local adaptation in Tsuga heterophylla habitat varied among years. In contrast, relative differences in flowering time among seed sources were stable across sites and years. Populations from the arid steppe were the earliest to flower; flowering was latest in populations from the mesic Tsuga heterophylla habitat. In terms of net reproductive rate, evidence for local adaptation in B. tectorum was obtained in populations from habitats representing environmental extremes: an arid, saline site dominated by the shrub Sarcobatus vermiculatus and clearings within the cool, mesic Tsuga heterophylla forest habitat. Unlike the plants introduced from other sites, members of the resident population at the Sarcobatus site flowered and produced seeds before soil water became limiting. In contrast, net reproductive rates in other habitats were sometimes the lowest for populations in their home site. This lack of an advantage for local populations within more environmentally moderate sites suggests that limited dispersal may restrict the rate at which superior genotypes are introduced into a particular site.     

Effects of competition with Bromus tectorum on early establishment of Poa secunda accessions: can seed source impact restoration success?

When landscapes are heavily impacted by biological invasion, local populations of native plant species may no longer be adapted to altered environmental conditions. In these cases, it is useful to investigate alternative sources of germplasm, such as cultivated varieties, for planting at restoration sites. This study compared cultivated and wild (local) varieties of the native perennial bunchgrass, Poa secunda J. Presl, grown with and without the exotic, invasive Bromus tectorum L. in a greenhouse setting. While P. secunda cultivars emerged and grew more rapidly than wild seed sources, this advantage declined in the presence of B. tectorum and cultivated germplasm did not outperform wild accessions in the presence of an invasive species. Given the novel genetic background of cultivars and their potential to alter patterns of dominance in native plant communities, we recommend the use of local or regional wild seed sources when possible to conserve regional patterns of genetic diversity and adaptation. Use of multiple seed sources may increase the potential for capturing vigorous genotypes in the restoration seed mix. In cases where sites are heavily impacted by exotic, invasive species, other control measures will be necessary to improve establishment of native species in grassland restoration programs.     

Consequences of Seed Origin and Biological Invasion for Early Establishment in Restoration of a North American Grass Species

Local, wild-collected seeds of native plants are recommended for use in ecological restoration to maintain patterns of adaptive variation. However, some environments are so drastically altered by exotic, invasive weeds that original environmental conditions may no longer exist. Under these circumstances, cultivated varieties selected for improved germination and vigor may have a competitive advantage at highly disturbed sites. This study investigated differences in early establishment and seedling performance between wild and cultivated seed sources of the native grass, Poa secunda, both with and without competition from the invasive exotic grass, Bromus tectorum. We measured seedling survival and above-ground biomass at two experimental sites in western Montana, and found that the source of seeds selected for restoration can influence establishment at the restoration site. Cultivars had an overall advantage when compared with local genotypes, supporting evidence of greater vigor among cultivated varieties of native species. This advantage, however, declined rapidly in the presence of B. tectorum and most accessions were not significantly different for growth and survival in competition plots. Only one cultivar had a consistent advantage despite a strong decline in its performance when competing with invasive plants. As a result, cultivated varieties did not meet expectations for greater establishment and persistence relative to local genotypes in the presence of invasive, exotic species. We recommend the use of representative local or regional wild seed sources in restoration to minimize commercial selection, and a mix of individual accessions (wild, or cultivated when necessary) in highly invaded settings to capture vigorous genotypes and increase the odds native plants will establish at restoration sites.     

Studies on the microbial populations of the rhizosphere of big sagebrush (<Emphasis Type="Italic">Artemisia tridentata</Emphasis>)

Prolonged use of broad-spectrum antibiotics has led to the emergence of drug-resistant pathogens, both in medicine and in agriculture. New threats such as biological warfare have increased the need for novel and efficacious antimicrobial agents. Natural habitats not previously examined as sources of novel antibiotic-producing microorganisms still exist. One such habitat is the rhizosphere of desert shrubs. Here, we show that one desert shrub habitat, the rhizosphere of desert big sagebrush (Artemisia tridentata) is a source of actinomycetes capable of producing an extensive array of antifungal metabolites. Culturable microbial populations from both the sagebrush rhizosphere and nearby bulk soils from three different sites were enumerated and compared, using traditional plate-count techniques and antibiotic activity bioassays. There were no statistical differences between the relative numbers of culturable non-actinomycete eubacteria, actinomycetes and fungi in the rhizosphere versus bulk soils, but PCR amplification of the 16S rRNA gene sequences of the total soil DNA and denaturing gradient gel electrophoresis showed that the community structure was different between the rhizosphere and the bulk soils. A high percentage of actinomycetes produced antimicrobials; and the percentage of active producers was significantly higher among the rhizosphere isolates, as compared with the bulk soil isolates. Also, the rhizosphere strains were more active in the production of antifungal compounds than antibacterial compounds. 16S rRNA gene sequence analysis showed that sagebrush rhizospheres contained a variety of Streptomyces species possessing broad spectrum antifungal activity. Scanning electron microscopy studies of sagebrush root colonization by one of the novel sagebrush rhizosphere isolates, Streptomyces sp. strain RG, showed that it aggressively colonized young sagebrush roots, whereas another plant rhizosphere-colonizing strain, S. lydicus WYEC108, not originally isolated from sagebrush, was a poor colonizer of the roots of this plant, as were two other Streptomyces isolates from forest soil. These results support the hypothesis that the rhizosphere of desert big sagebrush is a promising source of habitat-adapted actinomycetes, producing antifungal antibiotics.     

VARIABILITY AMONG DANTHONIA SERICEA (GRAMINEAE) POPULATIONS IN RESPONSES TO SUBSTRATE MOISTURE LEVELS

Populations of Danthonia sericea from the New Jersey Pine Barrens were investigated as to possible ecological differentiation in moisture tolerances at germination, seedling, and mature plant stages. Field studies had indicated that populations with pubescent lemmas and leaf sheaths are restricted to well-drained sandy upland sites and that relatively glabrous populations are found in open bogs or low wet areas bordering rivers or ponds. Greenhouse studies of responses of clonal and seed materials to saturated, moist, and dry moisture levels showed significant differences among populations in germination, growth, and survival. Reciprocal transplants at upland and bog sites provided a field evaluation (1969–1974) of the survival and performance of the morphological variants in each other's habitat under competitive conditions. Restriction to respective habitats was found to be primarily intolerance of wet site factors for the pubescent plants coupled with an inability of glabrous plants to successfully compete in upland sites.     

荒漠草原-灌丛镶嵌体的植被稳态转变特征

以柠条锦鸡儿（Caragana korshinskii）灌丛引入形成荒漠草原-灌丛地镶嵌体为研究对象,选择其内部荒漠草地（DG）、草地边缘（GE）、灌丛边缘（SE）、灌丛地（SL）为试验样地,开展荒漠草原向灌丛地人为稳态转变过程土壤水分与植被变化特征研究。结果如下：各样地0-200 cm土层水分含量随着转变过程呈显著降低趋势（P<0.05）,其中荒漠草地、草地边缘和灌丛边缘样地的土壤水分均在秋季雨水补充期增加,灌丛地由于深层土壤水分过度消耗而愈加降低,且灌丛边缘和灌丛地深层水分显著低于荒漠草地、草地边缘（P<0.05）,垂直动态不显著;地上植被随样地转变除优势度指数表现为灌丛地多样性、均匀度、丰富度指数均显著低于各样地（P<0.05）,多年生优势草本蒙古冰草（Agropyron mongolicum）、短花针茅（Stipa breviflora）逐渐被一年生草本刺藜（Dysphania aristata）、猪毛蒿（Artemisia scoparia）所代替;土壤种子库萌发种类随样地转变逐渐减少,种子库密度则显著升高（P<0.05）,灌丛地以一年生草本植物为主（占总密度的97.51%）,荒漠草地则以多年生草本萌发为主,且各样地土壤萌发种子库及多年生草本多集中于0-5 cm土层（P<0.05）;转变过程荒漠草原和灌丛地植被地上地下相似度分别为0.14和0.19,均显著低于两边缘样地0.35,较两边缘样地群落更为稳定,呈现草原灌丛化过程中草地-灌丛共存的植被双稳态特征。     

Deposition of exotic bird-dispersed seeds into three habitats of a fragmented landscape in the northeastern United States

Dispersal by frugivorous birds facilitates invasion by many exotic plants. We measured the seed rain of ornithochorous native and exotic plants at three habitats of a fragmented landscape of the northeastern United States for 1 year. We studied maple-beech forests, old fields, and abandoned conifer plantations. Across all sites we collected 2,196 ornithochorous seeds, including seeds from six exotic species and 10 native species. The majority (90%) of collected seeds were from exotic species. Seed dispersal was broadly similar among habitats, though seed rain of exotic species was higher in old fields than forested habitats. Seed rain was not strongly influenced by artificial perches for most species. However, seeds of exotic species were more commonly found in traps under an artificial perch in old fields. Seed rains for the exotic Elaeagnus umbellata, Rhamnus cathartica, and Rosa multiflora were positively associated with local density of mature plants. Seed rain of R. cathartica was positively associated with abundance of seedlings but not saplings, suggesting that post-dispersal mortality was important. Seed dispersal of the exotic Lonicera spp. was high in all habitats, accounting for 66% of all seeds collected. With the exception of Lonicera spp., seed rain of common exotic invaders was affected by the abundance of seed sources, and these species might be effectively controlled by elimination of local fruiting plants. Fruits of Lonicera morrowii, which has extensively invaded our area, are apparently a common component in the diet of frugivores.     

Experimental evidence for an alkali ecotype of <Emphasis Type="Italic">Lolium multiflorum</Emphasis>, an exotic invasive annual grass in the Central Valley,CA, USA

One of the characteristics of highly invaded ecosystems is that exotic species are often poor invaders of edaphically severe sites, which become refuges for native flora. To investigate the invasive potential of Lolium multiflorum (Per.) into alkali sites in California, an ex-situ reciprocal transfer experiment was carried out using seeds from populations of L. multiflorum taken from three sites differing in alkalinity (and inundation), including alkali sink soils (pH 8.5) and sink matrix soils (pH 7.4) located within meters of each other, and non-sink soils (pH 5.0) located several km away. Survivorship, plant height, leaf number and seed production were assessed. In addition, a native composite, Hemizonia pungens (Hick.), commonly found on alkali sinks was also sampled at the sink and sink matrix microsites. Lolium multiflorum plants grown from alkali sink and sink matrix seeds produced fewer leaves and seeds but were taller than plants grown from non-alkali seeds, the latter perhaps an adaptation to frequently inundated soils. Non-alkali genotypes fared poorly in sink soils for all traits, both in comparison to their growth on non-sink soils, and in comparison to the sink and sink edge genotypes. This suggests the existence of L. multiflorum ecotypes adapted to inundated alkali sinks, a genotypic difference that occurs on a broad spatial scale (kilometers), but not so obviously on the micro-site scale (meters) between sink and sink matrix populations. These data suggest that the absence of exotic invasives from alkali sites may be temporary if they are evolving tolerance for these severe sites, and this may threaten the future of the native alkali specialists that currently find refuge in these sites.     

Drivers of <Emphasis Type="Italic">Bromus tectorum</Emphasis> Abundance in the Western North American Sagebrush Steppe

Bromus tectorum can transform ecosystems causing negative impacts on the ecological and economic values of sagebrush steppe of the western USA. Although our knowledge of the drivers of the regional distribution of B. tectorum has improved, we have yet to determine the relative importance of climate and local factors causing B. tectorum abundance and impact. To address this, we sampled 555 sites distributed geographically and ecologically throughout the sagebrush steppe. We recorded the canopy cover of B. tectorum, as well as local substrate and vegetation characteristics. Boosted regression tree modeling revealed that climate strongly limits the transformative ability of B. tectorum to a portion of the sagebrush steppe with dry summers (that is, July precipitation <10 mm and the driest annual quarter associated with a mean temperature >15°C) and low native grass canopy cover. This portion includes the Bonneville, Columbia, Lahontan, and lower Snake River basins. These areas are likely to require extreme efforts to reverse B. tectorum transformation. Our predictions, using future climate conditions, suggest that the transformative ability of B. tectorum may not expand geographically and could remain within the same climatically suitable basins. We found B. tectorum in locally disturbed areas within or adjacent to all of our sample sites, but not necessarily within sagebrush steppe vegetation. Conversion of the sagebrush steppe by B. tectorum, therefore, is more likely to occur outside the confines of its current climatically optimal region because of site-specific disturbances, including invasive species control efforts and sagebrush steppe mismanagement, rather than climate change.     

Seed harvesting by a generalist consumer is context‐dependent: Interactive effects across multiple spatial scales

Granivore foraging decisions affect consumer success and determine the quantity and spatial pattern of seed survival. These decisions are influenced by environmental variation at spatial scales ranging from landscapes to local foraging patches. In a field experiment, the effects of seed patch variation across three spatial scales on seed removal by western harvester ants Pogonomyrmex occidentalis were evaluated. At the largest scale we assessed harvesting in different plant communities, at the intermediate scale we assessed harvesting at different distances from ant mounds, and at the smallest scale we assessed the effects of interactions among seed species in local seed neighborhoods on seed harvesting (i.e. resource–consumer interface). Selected seed species were presented alone (monospecific treatment) and in mixture with Bromus tectorum (cheatgrass; mixture treatment) at four distances from P. occidentalis mounds in adjacent intact sagebrush and non‐native cheatgrass‐dominated communities in the Great Basin, Utah, USA. Seed species differed in harvest, with B. tectorum being least preferred. Large and intermediate scale variation influenced harvest. More seeds were harvested in sagebrush than in cheatgrass‐dominated communities (largest scale), and the quantity of seed harvested varied with distance from mounds (intermediate‐scale), although the form of the distance effect differed between plant communities. At the smallest scale, seed neighborhood affected harvest, but the patterns differed among seed species considered. Ants harvested fewer seeds from mixed‐seed neighborhoods than from monospecific neighborhoods, suggesting context dependence and potential associational resistance. Further, the effects of plant community and distance from mound on seed harvest in mixtures differed from their effects in monospecific treatments. Beyond the local seed neighborhood, selection of seed resources is better understood by simultaneously evaluating removal at multiple scales. Associational effects provide a useful theoretical basis for better understanding harvester ant foraging decisions. These results demonstrate the importance of ecological context for seed removal, which has implications for seed pools, plant populations and communities.     

Subtle topographical differences along a floodplain promote different plant strategies among Paspalum dilatatum subspecies and populations

It was hypothesised that subtle topographical differences might cause the existence of ecotypes along a floodplain. The apomict grass Paspalum dilatatum subspecies dilatatum inhabits flood‐prone lowlands as well as nearby uplands in the floodplains of Argentina, while the sexual P. dilatatum subspecies flavescens almost exclusively inhabits the uplands. The aim of the present study was to identify the different traits that allow these P. dilatatum populations to inhabit different habitats. Plants of P. dilatatum were reciprocally transplanted between uplands and lowlands. Morphophysiological traits related to flooding tolerance were measured during a flood. Subspecies dilatatum from the uplands and subspecies flavescens showed a high physiological performance in the uplands but a considerable decrease in stomatal conductance, net photosynthesis rates and tiller number in the flooded lowlands. In contrast, the subspecies dilatatum from the lowlands showed relatively lower and stable stomatal conductance, photosynthesis rates and leaf water potential at both sites. Subspecies dilatatum from the lowlands outperformed upland populations at the lowland site with respect to tillering. Leaves of subspecies dilatatum from the lowlands that had grown at the lowland habitat had a lower blade/sheath proportion than leaves of plants transplanted to the uplands. This behavior did not occur in both upland populations. Results suggest that dilatatum Lowland plants have the typical strategy of stress‐tolerant genotypes and that the upland populations are adapted to habitats where competitive species are selected. In conclusion, habitats with subtle differences in topographic level can favour both ecotypic differentiations within an apomict subspecies but also the maintenance of morphophysiological similitudes between coexisting upland populations belonging to different subspecies.     

Contingent fire effects on granivore removal of exotic woody plant seeds in longleaf pine savannas

Prescribed fire is increasingly used to inhibit woody encroachment into fire-dependent ecosystems, yet its effects on other processes influencing invasion are poorly understood. We investigated how fire influences exotic woody invasion through its effects on granivore activity, and whether these effects depend on the habitat in which seed predation takes place. We quantified seed removal for four species of exotic woody plants (Albizia julibrissin, Elaeagnus umbellata, Melia azedarach and Triadica sebifera) in 17 sites in longleaf pine savanna that varied in time since fire (one or three growing seasons post-fire) in the sandhills region of North Carolina, USA. Within each site, we established paired plots in upland and upland-wetland ecotone communities and presented seeds in depots that allowed either arthropod, or arthropod and small vertebrate access. We found that differences in seed removal with time since fire were contingent on habitat and granivore community. In ecotones, three of four species had higher proportions of seeds removed from plots that were three growing seasons post-fire than plots one growing season post-fire, whereas only T. sebifera showed this pattern in upland habitat. Allowing vertebrate granivores access to seeds enhanced seed removal, and this effect was strongest in ecotone habitat. While granivores removed seeds of all four plant species, removal of E. umbellata was significantly higher than that of the other species, suggesting that granivores exhibited seed selection. These findings suggest that ecotone habitats in this system experience greater seed removal than upland habitats, particularly as time since fire increases, and differences are mainly due to the activity of vertebrate granivores. Such differences in seed removal, together with seed selection, may contribute to variation in exotic woody invasion of longleaf pine savannas.     

Processes regulating the invasion of European buckthorn (Rhamnus cathartica) in three habitats of the northeastern United States

Invasion by exotic plants often is restricted by processes, such as seed predation, acting on early life-history stages; however, the relative importance of these processes might vary among habitats. Modern human land use has created a mosaic of habitats in many landscapes, including the landscape of the northeastern United States. European buckthorn (Rhamnus cathartica) is an exotic plant that has achieved varying success in North American habitats. We studied dispersal, seed survival, germination, and seedling survival of buckthorn populations at four plots in each of sugar maple (Acer saccharum) forests, old fields, and abandoned conifer plantations in central New York State. Dispersal was low in maple forests, as evidenced by low collection rates of R. cathartica seeds in seed traps. Rates of post-dispersal seed predation were highest in maple forests and lowest in old fields, suggesting greater use of maple forests by granivorous rodents. Germination rates did not vary among seeds planted in soils of these habitats when studied in the laboratory despite differences in soil pH. Survival of transplanted seedlings was low in maple forests relative to old fields and plantations. Buckthorn invasion of old fields and abandoned plantations was not strongly constrained by factors we considered, and the buckthorn populations in these habitats were large. A combination of low dispersal by frugivores, low seed survival due to predation, and low seedling survival due to dim light conditions apparently prevents R. cathartica from invasion of intact maple forests of our area. Native fauna and canopy closure may act synergistically to reduce success of invasive plants in natural habitats.     

人工柠条-荒漠草地交错带拟步甲昆虫群落多样性

选取宁夏东部盐池县人工柠条灌丛-荒漠草地交错带为研究样地,以距离梯度形式,利用巴氏诱罐法对拟步甲科昆虫群落多样性进行了调查,研究了边缘效应对拟步甲昆虫群落多样性的影响。共采集拟步甲昆虫1405只,分属8属13种。结果表明,克小鳖甲和弯齿琵甲个体数量分别占总个体数的32.74%和43.27%,为该地区的优势种类。拟步甲昆虫在人工柠条灌丛-荒漠草地交错带的分布可分为3种类型,优势种克小鳖甲和弯齿琵甲属于栖息地广布型,但二者并没有表现出十分均匀的适应性,而且他们在各样带的分布呈显著负相关,克小鳖甲的分布趋向于沙生荒漠环境,弯齿琵甲的分布趋向于柠条灌丛;常见种蒙古漠王、小皮鳖甲的分布偏向于喜好荒漠草地;异距琵甲、奥氏真土甲、网目土甲和淡红毛隐甲的分布则偏向于喜好柠条灌丛。拟步甲昆虫丰富度与Shannon-Wiener多样性沿边缘分别向柠条灌丛和荒漠草地内部降低,个体数量总体上沿柠条灌丛、交错带向荒漠草地内部降低,但各样带差异不显著。从季节动态看,荒漠草地和柠条灌丛物种多样性季节变化相似,3种生境拟步甲活动密度季节变化相似。边缘效应强度分析呈现边缘正效应。CCA排序图显示分类地位相近的物种在生态适应上具有趋同性。多元回归分析表明,植被密度影响拟步甲昆虫群落的优势种和个体数量,植被盖度和高度分别影响拟步甲群落的均匀度和丰富度。主坐标分析(PCo A)排序表明人工柠条灌丛-荒漠草地交错带的拟步甲昆虫群落组成与柠条灌丛内部没有明显分化,但趋于向柠条灌丛生境演替。     

The importance of facilitation in the zonation of shrubs along a coastal salinity gradient

Question: What are the interactive roles of abiotic stress and plant interactions in mediating the zonation of the shrub Tamarix chinensis along a salinity gradient? Location: Yellow River estuary (37°46′N, 119°09′E), northeast China. Methods: We surveyed the zonation of T. chinensis along a salinity gradient and quantified its salt tolerance using a pot experiment. In two field experiments, we transplanted T. chinensis seedlings into salt marsh, transitional zone and upland habitats, manipulated neighbours and quantified survivorship and biomass to examine neighbour effects. We also quantified vegetation effects on abiotic conditions in each zone. Results: Tamarix chinensis dominated the transitional zone, but was absent in upland and salt marsh habitats. In the pot experiment, T. chinensis grew well in freshwater treatments, but was inhibited by increasing salinity. Field experiments revealed that competition from neighbours limited T. chinensis growth in the uplands, while T. chinensis transplants were limited, with or without neighbours, in the salt marsh by high soil salinity. In the transitional zone, however, T. chinensis transplants performed better with than without neighbours. Vegetation removal significantly elevated soil salinity in the transitional zone, but not in other zones. Conclusions: Competition, facilitation and abiotic stress are all important in mediating the zonation of T. chinensis. Within its physiological stress tolerance range, or fundamental niche, it is limited by plant competition in low salinity habitats, and facilitated by neighbours in high salt stress habitats, but cannot survive in salt marshes having salinities above its salt stress tolerance limit. Our results have implications for understanding the relationships between facilitation and stress gradients.