Predicting the spread of an invasive plant: combining experiments and ecological niche model

Rapid evolutionary adjustments to novel environments may contribute to the successful spread of invasive species, and can lead to niche shifts making range dynamics unpredictable. These effects might be intensified by artificial selection in the course of breeding efforts, since many successful plant invaders were deliberately introduced and cultivated as ornamentals. We hypothesized that the invasion success of Buddleja davidii, the ornamental butterfly bush, is facilitated by local adaptation to minimum temperatures and thus, exhibits unpredictable range dynamics. To assess the potential effects of adaptive evolution and artificial selection on the spread of B. davidii, we combined a common garden experiment investigating local adaptation to frost, with ecological niche modelling of the species’ native and invasive ranges. We expected that populations naturalized in sub‐continental climate are less susceptible to frost than populations from oceanic climate, and that the invasive range does not match predictions based on climatic data from the native range. Indeed, we revealed significant variation among invasive B. davidii populations in frost resistance. However, frost hardiness was not related to geographic location or climatic variables of the populations’ home site, suggesting that invasive B. davidii populations are not locally adapted to minimum temperatures. This is in line with results of our ecological niche model that did not detect a niche shift between the species’ native range in China, and its invasive range in Europe and North America. Furthermore, our niche model showed that the potential invasive range of B. davidii is still not completely occupied. Together with the frost resistance data obtained in our experiment, the results indicate that climatic conditions are currently not limiting the further spread of the species in Europe and North America.     

A century of the ornamental plant trade and its impact on invasion success

We identify a significant relationship between domestic market-based propagule pressure, as measured both in presence in the British horticultural market and in seed prices of ornamental plant species, with success in invasion. We employ a multispecies temporal approach and use a Generalized Estimation Equation model comparing ornamental non-native species introduced into Britain which started to invade with species introduced but not known outside cultivation. Historical nursery catalogues gave information on the availability and prices of seeds of 506 ornamental species in the British horticultural market every 20 years from 1885 to 1985. Higher market frequency and cheap prices of seeds were more significant and had a greater impact on the invading probability 20 years later than at the date of listing in a nursery catalogue. Our results suggest that national economic factors are an important part of the explanation for the invasiveness of ornamental plant species, and hence for the development of potential solutions.     

Hardiness dependent accumulation of phospholipids in leaves of wheat cultivars

Phospholipid content and composition in the leaves of thirteen cultivars of wheat differing in frost hardiness, were compared before and after hardening in order to see whether phospholipids play a role in surviving at low temperatures. The content of phospholipids in the leaves at the end of the hardening could be related to the aquired hardiness, and the relationship between the two phenomena could be described by the regression equation y = (0.174 ± 0.0016) x + 1311. Accumulation of phospholipids represents a hardiness dependent augmentation of membranes. Phosphatidylcholine appeared to be preferentially accumulated in hardy cultivars during the hardening process. As there are no significant differences in phospholipid fatty acid compositions investigated earlier, these results suggest the importance of the polar head group composition of membranes in avoiding frost injury.     

Changes in fatty acid composition of winter wheat during frost hardening

Twelve day old winter wheat seedlings (cvs Kharkov, frost hardy and Champlein, less hardy) accumulated linolenic acid at the expense of linoleic acid during controlled hardening. The change was most pronounced in the roots, where it was not specific to the phospholipid fraction. It was less marked in the leaves, but occurred there mainly in the phospholipids. The lack of differences between fatty acid profiles of the two cultivars rules out the explanation of varietal differences in frost hardiness in winter wheat on the basis of major changes in fatty acid unsaturation.     

Characteristics of the rRNA synthesis taking place at low temperatures in wheat cultivars with varying degrees of frost hardiness

The characteristics and intensity of rRNA synthesis occurring at temperatures around freezing point were examined in connection with frost hardiness in wheat cultivars. The intensity of rRNA synthesis taking place near freezing point showed a close positive correlation with the critical temperature (LT₅₀) characteristic of the frost resistance of the varieties. The quantity of high MW precursors to rRNA appears to be heterogeneous in hardy and non hardy varieties at low temperatures and point primarily to an inhibition in the maturation processes of ribosomes in the final nuclease processing step in non-frosthardy varieties.     

Genetic relationships among three native North-American Mahonia species, invasive Mahonia populations from Europe, and commercial cultivars

Horticulture is one of the most important pathways for plant invasion. We used microsatellite markers to reveal the impact of plant breeding on Mahonia aquifolium, an invasive ornamental shrub. Since it was bred by hybridization with the related species M. repens and M. pinnata, we compared populations of the three native species, various commercial cultivars and invasive populations. Invasive populations and cultivars were genetically differentiated from the native groups, but differences did not result from genetic bottlenecks. In cultivars but not in invasive populations, we proved genes from M. pinnata. No significant amount of M. repens genes were found in cultivars and invasive populations, but this result has to be viewed with caution because of the close relationship between native M. aquifolium and M. repens. We conclude that the evolution of invasive Mahonia populations was a result of restriction of gene pool during introduction, secondary release, and artificial selection, in combination with an increase of genetic diversity by plant breeders and by extensive gene flow.     

Accumulation of Dehydrins and ABA-Inducible Proteins in Wheat Seedlings during Low-Temperature Acclimation

Using one-dimensional SDS-PAGE and immunochemical methods, we detected the presence and estimated the content of dehydrins and ABA-inducible (RAB) polypeptides in etiolated seedlings of four spring and three winter wheat (Triticum aestivum L.) cultivars differing in frost hardiness. We hardened three-day-old seedlings at 4°C for nine days or grew them at 22°C for a day (control seedlings). We established that heat-stable cold-regulated (COR) polypeptides with mol wts of 209, 196, 169, 66, 50, and 41 kD, which are characteristic of hardened wheat seedlings, were homologous to polypeptides from a dehydrin family and polypeptides with mol wts of 209, 196, 66, 50, and 41 kD were immunologically related to RAB-proteins. We supposed that these COR polypeptides were involved in the prevention of local protein dehydration and denaturation during hypothermia. Analysis of the relative content of COR proteins revealed a close correlation between the cultivar frost hardiness and the concentration of these proteins. It seems evident that different accumulation of dehydrins and RAB polypeptides in different cultivars of a single species is one of the causes for different plant frost hardiness.     

Are plant populations in expanding ranges made up of escaped cultivars? The case of Ilex aquifolium in Denmark

Rapid range expansions are becoming more prevalent, especially as climate continues to change. The escape of ornamental plants within their native range represents a significant, but often overlooked component of this process. Few studies have focused on the role of ornamental plants in range expansions using molecular markers to identify the possibility of mixed native and cultivar populations. The purpose of the present study was to determine the genetic variation of a native woody plant with ornamental conspecifics at the edge of its distribution. We selected Ilex aquifolium L. (English holly) which grows naturally in Denmark but is spreading eastward in Scandinavia, possibly due to a combination of climate change and introduction of more frost-tolerant cultivars. We sampled 187 individuals from older and recently established populations, and cultivated I. aquifolium throughout Denmark, and compared them using nuclear SSR and AFLP. The overall results showed no structure or clustering of plants from the historical or the expanding range, or of wild or cultivated plants. The only clusters found were for a group of cultivated hybrid I. aquifolium. The samples represented four genetic groupings, indicating either hybridization between cultivated and wild I. aquifolium or replacement of the latter by cultivars. Thus, ornamental genotypes of I. aquifolium contribute to the northeastern range expansion of the species and eventual invasiveness within its native range.     

Winter hardiness in trench celery

A total of 51 red and white commercial varieties of trench celery were grown and examined for winter hardiness at the National Vegetable Research Station. Although some plants of most varieties survived the winter, none showed a commercially useful degree of winter hardiness, in that none were marketable after moderate frost. The red varieties had a significantly better survival than the white varieties. In the first season, single plant selections were made from the varieties having both the highest percentage of winter survival and the best agronomic characters. Although some of the progenies derived from these selected plants by self-pollination showed a small increase in winter survival as compared with their parent varieties when both were grown in the second season, no plants of these progenies were sufficiently hardy to remain marketable after moderate frost. It was concluded that none of the varieties tested showed any promise as a source of winter hardiness to be used in a breeding programme, and that selection for one generation from within these varieties did not improve winter hardiness to a useful extent.     

Niche based distribution modelling of an invasive alien plant: effects of population status, propagule pressure and invasion history

Forecasting the spatial spread of invasive species is important to inform management planning. Niche-based species distribution models offer a well-developed framework for assessing the potential range of species. However, these models assume equilibrium between the species’ distribution and its ecological requirements. During range expansion, invasive species are not in such equilibrium due to both dispersal limitation and frequent casual occurrence in sites unsuitable to persistent populations. In this article we use the example of the invasive annual plant Ambrosia artemisiifolia in Austria to evaluate if model accuracy can be enhanced in such non-equilibrium situations by taking account of propagule pressure and by restricting model calibration to naturalized populations. Moreover, we test if model accuracy increases during invasion history using distribution data from 1984 to 2005. The results suggest that models calibrated with naturalized populations are much more accurate than those based on the total set of records. Proxies of propagule pressure slightly but significantly improve goodness of fit, accuracy, and Type I and II error rates of models calibrated with all available records but have less consistent effects on models of naturalized populations. Model accuracy did not increase during the recent invasion history, probably because the species is still far from an equilibrium distribution. We conclude that even a coarse assessment of population status with records of invasive species delivers important information for predictive modelling and that proxies of propagule pressure should be included into such models at least during early to intermediate stages of the invasion history.     

Revealing Hereditary Variation of Winter Hardiness in Cereals

A set of cereal crops and differentiating cultivars was shown to be of utility for identifying the major abiotic factors that limit the survival of winter crops in the cold season of a particular year. With this approach, the season was identified (1997–1998, Belgorod) when the survival of cereals depended on the tolerance to anaerobiosis rather than on the frost resistance. Differentiation of common wheat cultivars with respect to this property was attributed to a locus designated Win1 (Winter hardiness 1) and localized 3.2–5.8% recombination away from the B1 (awnlessness 1) gene. Winter barley (cultivar Odesskii 165) displayed the highest tolerance to anaerobiosis in the cold season; low and intermediate tolerance was established for winter durum wheat (cultivar Alyi Parus) and winter common wheat, respectively. Frost resistance and winter hardiness type 1 proved to be determined by different genetic systems, which showed no statistical association. Correlation analysis revealed significant positive associations of frost resistance in the field (1996–1997, Belgorod) with productivity, sedimentation index (Zeleny test), plant height, and vegetation period in wheat. Statistical analysis associated frost resistance with gliadin-coding alleles of homeologous chromosomes 1 and 6 of the A, B, and D wheat genomes.     

Saving camels from straws: how propagule pressure-based prevention policies can reduce the risk of biological invasion

Nonnative species that harm or have the potential to cause harm to the environment, economy, or human health are known as invasive species. Propagule pressure may be the most important factor in establishment success of nonnative species of various taxa in a variety of ecosystems worldwide, and strong evidence is emerging that propagule pressure determines both the scale of invasion extent and impact. In a limited way, the US government is applying a “propagule pressure approach” in a variety of prevention policy contexts aimed at minimizing the impact of harmful organisms. However, there are also readily apparent opportunities for enacting propagule pressure-based measures to fill current gaps in invasive species prevention and control at national, state, and local levels. An explicit focus on propagule pressure-based policies could substantially increase the effectiveness of US efforts to prevent the introduction of invasive species through by intentional and unintentional introductions. The views expressed in this paper are solely those of the authors and do not necessarily reflect those of the US government. “As the last straw breaks the laden camel’s back...” -Charles Dickens, Dombey and Son     

Effects of resource availability and propagule supply on native species recruitment in sagebrush ecosystems invaded by Bromus tectorum

Resource availability and propagule supply are major factors influencing establishment and persistence of both native and invasive species. Increased soil nitrogen (N) availability and high propagule inputs contribute to the ability of annual invasive grasses to dominate disturbed ecosystems. Nitrogen reduction through carbon (C) additions can potentially immobilize soil N and reduce the competitiveness of annual invasive grasses. Native perennial species are more tolerant of resource limiting conditions and may benefit if N reduction decreases the competitive advantage of annual invaders and if sufficient propagules are available for their establishment. Bromus tectorum, an exotic annual grass in the sagebrush steppe of western North America, is rapidly displacing native plant species and causing widespread changes in ecosystem processes. We tested whether nitrogen reduction would negatively affect B. tectorum while creating an opportunity for establishment of native perennial species. A C source, sucrose, was added to the soil, and then plots were seeded with different densities of both B. tectorum (0, 150, 300, 600, and 1,200 viable seeds m⁻²) and native species (0, 150, 300, and 600 viable seeds m⁻²). Adding sucrose had short-term (1 year) negative effects on available nitrogen and B. tectorum density, biomass and seed numbers, but did not increase establishment of native species. Increasing propagule availability increased both B. tectorum and native species establishment. Effects of B. tectorum on native species were density dependent and native establishment increased as B. tectorum propagule availability decreased. Survival of native seedlings was low indicating that recruitment is governed by the seedling stage.     

Non-native competitive perennial grass impedes the spread of an invasive annual grass

Invasive plants are degrading wildlands around the globe by displacing native species, reducing biodiversity, and altering ecological functions. The current approach of applying herbicides to invasive plants in wildlands has not been effective at curtailing their expansion and, in certain circumstances, may do more harm than good. Preventing the spread of invasive species has been identified as an important strategy to protect wildlands. However, few prevention strategies have actually been tested. We hypothesized that establishing competitive vegetation next to infestations would increase the biotic resistance of the plant community to invasion and decrease the invasive species propagule pressure beyond the competitive vegetation. To evaluate this, we established twelve competitive vegetation barriers in front of invasive annual grass, Taeniatherum caput-medusae (L.) Nevski, infestations. The non-native perennial grass Agropyron desertorum (Fisch. ex Link) Schult. was seeded into plant communities adjacent to the infestations to create the competitive vegetation barriers. Soil nutrient concentrations and the spread of T. caput-medusae were compared between where A. desertorum was seeded and not seeded (control treatment) 3 years after treatment. Less T. caput-medusae and lower soil ammonium and potassium concentrations in the competitive vegetation barrier than control treatment (P ≤ 0.01) suggest that establishing competitive vegetation increased the biotic resistance of the plant communities to invasion. Taeniatherum caput-medusae cover and density in the plant communities protected by the competitive vegetation barrier (locales across the barriers from the infestations) were ~42- and 47-fold less, respectively, than unprotected plant communities (P < 0.01). This suggests that invasive plant propagule pressure was decreased in the plant communities protected by competitive vegetation barriers. The establishment of competitive vegetation around infestations may be an effective strategy to prevent or at least reduce the spread of invasive plant species.     

Microclimate predicts frost hardiness of alpine Arabidopsis thaliana populations better than elevation

In mountain regions, topological differences on the microscale can strongly affect microclimate and may counteract the average effects of elevation, such as decreasing temperatures. While these interactions are well understood, their effect on plant adaptation is understudied. We investigated winter frost hardiness of Arabidopsis thaliana accessions originating from 13 sites along altitudinal gradients in the Southern Alps during three winters on an experimental field station on the Swabian Jura and compared levels of frost damage with the observed number of frost days and the lowest temperature in eight collection sites. We found that frost hardiness increased with elevation in a log‐linear fashion. This is consistent with adaptation to a higher frequency of frost conditions, but also indicates a decreasing rate of change in frost hardiness with increasing elevation. Moreover, the number of frost days measured with temperature loggers at the collection sites correlated much better with frost hardiness than the elevation of collection sites, suggesting that populations were adapted to their local microclimate. Notably, the variance in frost days across sites increased exponentially with elevation. Together, our results suggest that strong microclimate heterogeneity of high alpine environments can preserve functional genetic diversity among small populations. Synthesis: Here, we tested how plant populations differed in their adaptation to frost exposure along an elevation gradient and whether microsite temperatures improve the prediction of frost hardiness. We found that local temperatures, particularly the number of frost days, are a better predictor of the frost hardiness of plants than elevation. This reflects a substantial variance in frost frequency between sites at similar high elevations. We conclude that high mountain regions harbor microsites that differ in their local microclimate and thereby can preserve a high functional genetic diversity among them. Therefore, high mountain regions have the potential to function as a refugium in times of global change.     

Propagule pressure and environmental conditions interact to determine establishment success of an invasive plant species,glossy buckthorn (<Emphasis Type="Italic">Frangula alnus</Emphasis>), across five different wetland habitat types

Many invasive plant species are able to establish within a wide range of community types. This establishment success depends on high propagule pressure and successful recruitment of seedlings following propagule dispersal into receptive environments. This study aimed to investigate interactions between propagule pressure and environmental resistance to seedling recruitment of the invasive shrub, glossy buckthorn (Frangula alnus Mill.), over a range of wetland habitat types. We measured propagule deposition using seed traps and recruitment success using sown plots, while characterizing vegetation and abiotic environmental conditions in five adjacent wetland habitat types. Drier habitats, which included Cedar Swamp, Shrub Carr, and String, had lower resistance to buckthorn establishment than the wetter Flark and Cattail Marsh. The drier habitats supported more woody species and provided more raised hummock surfaces essential for successful buckthorn recruitment and establishment. Propagule pressure was also higher in dry habitats that supported higher densities of adult glossy buckthorn, while long-distance dispersal into areas with low adult density was uncommon. Natural recruitment was highest in sites with intense propagule pressure, but experimental sowing of seeds demonstrated that buckthorn establishes in wet sites with higher resistance if propagule pressure is increased and seeds are deposited on hummocks. This study demonstrates the affinity of glossy buckthorn for drier wetland sites, and provides empirical evidence that environmental resistance can be overcome by higher propagule pressure.     

Genetic analysis of frost hardiness traits in tuber-bearingSolanum species

The inheritance of frost hardiness and cold acclimation potential traits was studied in three segregating populations derived from a cross betweenSolanum commersonii Dun. PI 243503 (cmm) andSolanum cardiophyllum Lindl., PI 184762 (cph), two parental genotypes with contrasting frost hardiness and cold acclimation potential. The levels of frost hardiness and cold acclimation potential were expressed as the LT₅₀, the temperature at which 50% of the cells in leaf discs were killed, as measured by the ion leakage method, following a controlled freeze test There was considerable variation in both frost hardiness and cold acclimation potential in all three segregating populations (F₁ F₁ xcmm, and F₁ xcph). Frost hardiness and cold acclimation potential were not correlated, suggesting that these two traits are under independent genetic control. The analysis of generation means indicated that the variation for both traits could be best explained by an additive-dominance model, with additive gene effects the most important Broad-sense heritability was 0.73 and 0.74 in the F₁ population, for frost hardiness and cold acclimation potential, respectively, and was 0.85 for either trait in the F₁ xcmm population, indicating that these two traits are highly inheritable. Our results suggest that it should be possible to incorporate the frost hardiness and cold acclimation traits from S.commersonii into cultivated potato species.     

A nursery Area for Sharks and Rays in Northeastern Brazil

Synopsis Elasmobranch nurseries are habitats where females give birth to their young and juveniles spend their early life history. Hypotheses concerning nurseries suggest that these provide the young a better source of food and protection against predation, however the degree of exposure to predatory risk in nursery areas vary. According to Branstetter hypothesis, slow growing species are either born at relatively large sizes or use protected nursery grounds, whereas faster growing species tend to rely more on growth rates than others factors. Primary nurseries are habitats where parturition occurs and in which the young live for a short time and secondary nurseries are habitats in which juveniles are found after leaving the primary nursery and before reaching maturity. Rhizoprionodon porosus and Carcharhinus acronotus use coastal waters off Cai?ara do Norte as both primary and secondary nurseries, while Carcharhinus limbatus makes use of the area as a seasonal primary nursery in late spring and early summer. C. falciformis apparently makes use of the end of inner continental shelf and the outer continental shelf as a primary nursery. Gymnura micrura, Rhinobatos percellens and Narcine brasiliensis makes use of coastal and turbid waters of region as both primary and secondary nurseries throughout the year, while Dasyatis guttata utilizes the area as a primary nursery. Aetobatus narinari and Rhinoptera bonasus, although caught in lesser number, appear to use the region as both primary and secondary nurseries and only primary nursery respectively. Dasyatis marianae (endemic from Northeastern Brazil) may use coastal sandbanks as a primary nursery, while Dasyatis Americana apparently exploits waters about 10 – 20 m in depth with rocky reef substrate only as secondary nursery. There was a measurable degree of temporal and spatial separation among sharks’ species, which may effectively create separate ecological niches, whereas among rays, with the exception of the stingray D. marianae, a spatial and temporal overlaps were observed. The Branstetter’s hypothesis was supported.     

Studies of Frost Hardiness in Woody Plants. II. Effect of Temperature on Hardening

The effect of temperature on hardening was studied at temperatures ranging from 0° to −20° using twigs of willow and poplar. In October and in late April when the twigs are not very frost hardy, hardening at 0° produced a considerable increase in their frost hardiness, although the effectiveness of hardening at 0° decreased with a decrease in the environmental temperature. In twigs which could withstand continuous freezing without injury, hardening at −3° to −5° was most effective in increasing the frost hardiness of the twigs. Below −20°, only negligible increase was observed either in frost hardiness or sugar content.

The rate of starch to sugar conversion differed remarkably in different twig tissues. The starch in xylem was more slowly converted to sugar than that in the cortex. The optimum temperature for converting starch into sugar during frost hardening was also found to be −3° to −5°. In addition, the greater the effectiveness of the hardening treatment, the greater the rate of conversion from starch to sugar. The frost hardiness of a twig is closely related to the sugar content of the twig, especially in the xylem.

     

Propagule pressure and native species richness effects drive invasibility in tropical dry forest seedling layers

Understanding the factors that encourage or inhibit plant invasions is vital to focusing limited invasive control efforts within areas where they are most practical and cost-effective. To extend the range of contexts in which invasibility is studied and aid the development of practical strategies to limit damaging plant invasions, we set out to test the relative importance of native species richness, native seedling density, and invasive propagule pressure, on the invasibility of artificial assemblages of naturally occurring tropical woody seedling communities. Our greenhouse mesocosms included a species pool of twelve trees and woody shrubs native to South Florida's tropical hardwood hammocks, and an increasingly prevalent noxious woody invader of this system, Ardisia elliptica. We found that invader propagule pressure was the single most important factor determining community invasibility. We also revealed a positive relationship between invasibility and native species richness in our polyculture mesocosms. Because A. elliptica biomass production significantly differed among different native monocultures and was not related to overyielding in native polycultures, we suggest that the effect of species richness on invasibility in this experiment was the result of sampling effects rather than a true effect of diversity.Three broad findings hold potential for application in preventing and controlling plant invasions, especially in the seedling layers of tropical dry forests: (1) effective invasive control efforts will likely benefit from measures to minimize propagule pressure; (2) managers might do well to prioritize invasive monitoring and removal efforts on the most diverse habitats within a management region; and (3) while more data are necessary to further understand our finding of a lack of association between productivity and invasibility, management regimes aimed at maximizing primary productivity might not increase invasibility, and in fact, strategies for controlling invasive plants via the management of ecosystem productivity may be ineffective.