No adaptation to altitude in the invasive plant Erigeron annuus in the Swiss Alps

The altitudinal distribution of plants is restricted by various environmental factors, with climatic conditions being one of the primary constraints. Here, we investigate what limits the altitudinal range of the introduced species Erigeron annuus in the Swiss Alps. We planted offspring of E. annuus plants originating from different altitudes into two common gardens, one located at an altitude representing the main area of distribution (400 m) and the other close to the current altitudinal limit of E. annuus in Switzerland (1000 m). In both common gardens all established plants survived and grew vigorously during the growing season. However, there was high winter mortality of seedlings at 1000 m. Furthermore, plant phenology was delayed and reproductive output was reduced at 1000 m, although the seeds produced were larger. The general lack of adaptation to altitude and only moderate levels of plasticity suggest that there is little potential for E. annuus to persist beyond its current altitudinal limit in the Swiss Alps. However, climate warming might promote the upward range expansion of E. annuus by reducing winter mortality and by increasing the chance of producing seeds within the growing season.     

The resurrection of Erigeron parryi (Asteraceae)

Erigeron parryi was described in 1890 from a single collection from southwest Montana. No additional collections were made during the following 90 years. Recent floristic treatments have synonomized E. parryi under E. ochroleucus. I used principal components and discriminant analyses of morphological characters to examine the relationship of E. parryi to its presumed closest relatives, E. ochroleucus, E. radicatus, and E. scribneri, in light of several populations discovered in the past 20 years. Erigeron parryi should be considered a distinct species. It can be readily distinguished from closely related species based on indument and caudex morphology. It is allopatric with E. ochroleucus and E. scribneri and ecologically separated from E. radicatus. Erigeron scribneri is intermediate between E. ochroleucus and E. radicatus and is probably best treated as a variety of one or the other of these species.     

Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system

The relative importance of seed, habitat and microhabitat limitation has rarely been investigated for invasive non-native species, although this is critical for their effective management and for predicting future range expansion. Rosa rugosa is an abundant non-native shrub in coastal habitats of NW Europe; it is common along the Baltic coast but more scarce in exposed dunes of the North Sea. To investigate whether invasion of exposed dunes is limited by seed, habitat and microhabitat limitation, seedling emergence and establishment were examined in a factorial sowing, transplant and disturbance experiment. Twenty plots were randomly placed in each of five dune habitats (white dune, Empetrum dune, grey dune, outer dune heath, inner dune heath), and studied over 2 years. Seedling emergence in control subplots was zero in all habitats, whereas 96% and 98% of the undisturbed and disturbed seeded subplots produced seedlings. Disturbance had a positive effect on emergence and subsequent survival in white dune, outer and inner dune heath. Seedling survival and growth, and growth of transplanted seedlings, were markedly lower in grey dune. These findings indicate that establishment of R. rugosa is seed-limited in coastal dune habitats, and that the species is able to establish in both active and fixed dunes once seeds have arrived. Although differential seedling emergence and growth indicate that habitats differ in their degree of invasion susceptibility by R. rugosa, the positive influence of small-scale disturbance suggests microhabitat limitation in some dune habitats as well. Dune management should aim to reduce seed production and dispersal of R. rugosa in near-natural sites, and anthropogenic changes of habitat dynamics should be prevented.     

Conyza sumatrensis (Retz.) E. H. Walker: A new record from Pakistan

Conyza sumatrensis (Retz.) E. H. Walker (Astereae: Asteraceae) has been recorded in Pakistan for the first time. The species is closely associated with C. bonariensis and C. canadensis. It differs from C. bonariensis in its smaller capitula with a small series of marginal and disc florets that lack a purplish-pink apex, but resembles it in disciform capitula. C. sumatrensis differs from C. canadensis in having five-lobed disc florets and lacking well-developed ray florets. The species is widely distributed in the Potohar region, particularly in Rawalpindi and Islamabad.     

Seed development in the rare cold desert sand dune shrub Eremosparton songoricum and a comparison with other papilionoid legumes

No study has yet been carried out on seed development in a cold desert sand dune papilionoid legume. Thus, our primary aims were to (i) monitor seed development in the cold desert sand dune species Eremosparton songoricum from the time of pollination to seed maturity, and (ii) compare seed development in this species with that in other species of papilionoid legumes. Fruit and seed size, mass and seed moisture content, and seed imbibition, germination, desiccation tolerance and water retention during development (pollination to seed maturity) were monitored in the papilionaceous shrub E. songoricum in the Gurbantunggut Desert of northwest China. The duration of seed development was 40 days. Seeds reached physiological maturity 28 days after pollination (DAP), at which time 58% of them germinated and they had developed desiccation tolerance. Seeds became impermeable 36–40 DAP, when their moisture content was about 10%. The final stage of maturation drying occurred via loss of water through the hilum. The developmental stages and their timing (DAP) in seeds of E. songoricum are generally similar to those reported for other papilionaceous legumes with a water‐impermeable seed coat (physical dormancy). In general, the developmental features of seeds with water‐impermeable coats at maturity do not appear to be specific to habitat or phylogeny.     

Seed size variation in the palm Euterpe edulis and the effects of seed predators on germination and seedling survival

Intraspecific variation in seed size is common in wild plant populations and has important consequences for the reproductive success of individual plants. Multiple, often conflicting evolutionary forces mediated by biotic as well as abiotic agents may maintain such a variation. In this paper we assessed seed size variation in a population of the threatened, commercially important palm Euterpe edulis in southeast Brazil. We investigated (i) how this variation affects the probability of attack by vertebrate and invertebrate post-dispersal seed predators, and (ii) if seed size influences the outcome of seeds damaged by beetles in terms of seed germination and early survival of seedlings. Euterpe edulis seeds varied in diameter from 8.3 to 14.1 mm. Neither insects nor rodents selected the seeds they preyed upon based on seed size. Seed germination and total, shoot and root biomasses of one-year seedlings were significantly and positively affected by seed size. Root biomass and seedling survival were negatively affected by seed damage caused by a scolytid beetle (Coccotrypes palmarum) whose adults bore into seeds to consume part of the endosperm, but do not oviposit on them. Seed size had a marginally significant effect on seedling survival. Therefore, if any advantage is accrued by E. edulis individuals producing large seeds, this is because of greater seed germination success and seedling vigor. If this is so, even a relatively narrow range of variation in seed size as observed in the E. edulis population studied may translate into differential success of individual plants.     

Facilitation and sand burial affect plant survival during restoration of a tropical coastal sand dune degraded by tourist cars

Coastal sand dunes support various ecosystem services, including storm protection and tourism. Restoration programs are often critical to preserve this ecosystem due to its fragility and high degree of degradation. Dune restoration still suffers from a general lack of knowledge of the ecological processes controlling tropical dune communities. We investigated if facilitation can increase restoration success by assisting plant survival in a Brazilian coastal dune degraded by buggies at the Environmental Protection Area of Jenipabu, RN, Brazil. We performed two field experiments on dune crest sites degraded by buggies. The first experiment tested how the presence of established vegetation and coconut mesh, mimicking soil‐stabilizing effect of vegetation, facilitates seedling survival and establishment of the early successional dominant dune plant Canavalia maritima (Fabaceae). The second experiment tested if coconut mesh and initial irrigation would allow the establishment of C. maritima outside the vegetation, using both seeds and transplants. We found that the presence of established vegetation positively affected seedling survival. Application of mesh positively affected microclimatic conditions and slightly increased survival, but only for seeds. Initial irrigation only had a small effect on transplanted seedlings survival. Nevertheless, there was extreme seedling mortality in the experiment in particular from sand burial. We conclude that restoration of these dune crests is very difficult. Facilitation by vegetation can reduce plant mortality by sand burial and desiccation, but successful restoration is likely to require a major effort involving a very high number of seedlings or seeds.     

Chemical Composition and Antimicrobial and Allelopathic Activity of Tunisian Conyza sumatrensis (Retz.) E.Walker Essential Oils

Conyza sumatrensis (Retz.) E.Walker (Asteraceae) is a spontaneous annual herb, fairly widespread throughout Tunisia, which has rarely been studied or valued in any sector. Essential oils were obtained by hydrodistillation of different parts (flower heads, leaves, stems, and roots) of C. sumatrensis plants, which were collected in autumn (November 2007) at the flowering stage in the area of Monastir, Tunisia. In total, 98 compounds, representing 88.1–99.3% of the oil composition, were identified by GC‐FID and GC/MS analyses. The root essential oil was distinguished by its high content in acetylenes (matricaria ester, 4 ; 74.3%), while those from flower heads and leaves were dominated by oxygenated sesquiterpenes (61.1 and 50.3%, resp.). The oils of C. sumatrensis from Tunisia belonged to a matricaria ester/caryophyllene oxide chemotype. All the oils were evaluated for antibacterial, antifungal, and allelopathic activities. The results indicate that the leaf oil exhibited significant in vitro antibacterial activity against Enterococcus faecalis, Staphylococcus aureus, and Proteus mirabilis and that the C. sumatrensis oils isolated from the aerial parts presented high mycelia‐growth inhibition of Candida albicans and the filamentous fungi tested. Moreover, the essential oils of the different plant parts inhibited the shoot and root growth of Raphanus sativus (radish) seedlings. Indeed, the inhibition of the hypocotyl growth varied from 28.6 to 90.1% and that of the radicle from 42.3 to 96.2%.     

Genotype–environment interactions affect flower and fruit herbivory and plant chemistry of Arabidopsis thaliana in a transplant experiment

Large differences exist in flower and fruit herbivory between dune and inland populations of plants of Arabidopsis thaliana (Brassicaceae). Two specialist weevils Ceutorhynchus atomus and C. contractus (Curculionidae) and their larvae are responsible for this pattern in herbivory. We test, by means of a reciprocal transplant experiment, whether these differences reflect environmental influences or genetic variation in plant defense level. All plants suffered more damage after being transplanted to the dune site than after being transplanted to the inland site. Plants of inland origin suffered more flower and fruit herbivory than plants of dune origin when grown at the dune transplant site, but differences were much smaller at the inland site. Both flower damage by adult weevils and fruit damage by their larvae were subject to significant genotype × environment interactions. The observed pattern in herbivory is a strong indication for local adaption of plant defense to the level of herbivory by Ceutorhynchus. In order to identify the mechanism of defense, a quantitative analysis of glucosinolates was performed on the seeds with HPLC. Highly significant differences were found in glucosinolate types and total concentration. These patterns were mainly determined by the origin of the plants (dune or inland) and by a genotype × environment interaction. Herbivory was not significantly correlated to the concentration of glucosinolates in seeds. We therefore analyzed the total metabolic composition of seeds, using NMR spectroscopy and multivariate data analysis. Major differences in chemical composition were found in the water–methanol fractions: more glucosinolate and sucrose in the dune and more fatty acids, lipids and sinapoylmalate in the inland populations. We discuss which of these chemical factors could explain the marked differences in damage between populations.     

The adaptive biological reproductive response of the spurge Euphorbia pithyusa subsp. pithyusa on coastal and mountain bioclimates

Plants of nine Euphorbia pithyusa subsp. pithyusa populations growing in Mallorca (Balearic Islands) under distinct bioclimatic conditions were chosen for the study, from coastal, with a semi-arid thermo-Mediterranean bioclimate, to mountain (1250 m), with both meso-Mediterranean and supra-Mediterranean bioclimate. The cyathia and fruits differed significantly in size between the populations, with the coastal populations producing smaller cyathia and fruits. Flowering and fruiting were prolonged and extended through different orders of the dichasia. However, the third-order dichasia produced, independently of the environmental conditions, the highest number of fruits of all the populations. Although seeds of all the populations showed comparable germination capacity at 20 °C, seeds from the highland populations were slower to germinate and exhibited prolonged dormancy, suggesting a direct relationship between the degree of dormancy and rainfall. These differences were significant (p ≤ 0.05) and independent of the yearly climatic variations for each of the populations studied. The reproductive strategies permitting E. pithyusa to have success in the colonization of zones with different bioclimates are based on possessing: a generalist pollination system and the capacity to adapt the length and intensity of the flowering and fruiting periods to climatic conditions and to vary the size and dormancy levels of seeds.     

A new species of Erigeron (Asteraceae) and its relatives in southwestern Utah

A narrow endemic of southwestern Utah,Erigeron flagellaris A. Gray var.trilobatus Maguire ex Cronq., is given specific status asErigeron proselyticus Nesom. It is closely related to two other rare endemics of the Zion National Park area,Erigeron religiosus Cronq. andErigeron sionis Cronq., but it does not share a similarly close relationship with typicalE. flagellaris. Erigeron proselyticus is diploid;E. sionis is tetraploid and probably agamospermic; three diploid populations and several triploid populations ofE. religiosus are known, including a population of triploids from the site of a paratype collection of this species. Variation in these triploids suggests the possibility of hybridization withErigeron divergens Torr. & Gray, or with an unknown species no longer occurring sympatrically withE. religiosus.     

Effect of Interspecific Competition and Herbivory on the Recruitment of an Invasive Alien Plant: Conyza sumatrensis

Most studies on the spread of a species have been carried out retrospectively. Conyza sumatrensis provides an exception; it has only been in Great Britain since 1984 and thereby provides the opportunity to follow an invasion from its early stages. Many different factors could lead to the exclusion of invading species from new habitats. Here we report results of a field experiment to determine the recruitment ability and population dynamics of C. sumatrensis within treatments of different levels of interspecific competition and herbivory over three years. Interspecific competition was manipulated using three cultivation treatments giving low, medium and high intensity competition. Factorial combinations of mollusc, insect and vertebrate (rabbit) herbivory were achieved using both chemical and physical exclusion techniques. We show that although C. sumatrensis recruitment occurs in areas where disturbance has caused low interspecific competition, maintenance of a population during secondary succession is unlikely. Effects of herbivory were dependent on the type of herbivory and the plant stage grazed. Rabbit grazing had strong negative effects on recruitment and survival of C. sumatrensis throughout the experiment. Insect herbivory, however, had no effect on any life stage. Mollusc herbivory showed a significant interaction with interspecific competitors, reducing recruitment only where competition was experimentally reduced. This revised version was published online in July 2006 with corrections to the Cover Date.     

Functions of canopy-stored seeds in the dune ecosystem: conclusions from Agriophyllum squarrosum and Artemisia wudanica

Delay in seed release is found in the dune ecosystem. However, its functions have been rarely reported. We studied delayed seed release in the Agriophyllum squarrosum (Chenopodiaceae), an annual psammophyte, and the Artemisia wudanica (Asteraceae), a psammophilous subshrub in an active sand dune field in Inner Mongolia, China, to get insights into its functions in dealing with sand movement. We concluded that delay in seed release in the dune ecosystem are as follows: (1) postponing dispersal of a portion of seeds until the end of windy season and the start of growing season; (2) regulating the spatiotemporal pattern of seed bank; (3) helping psammophytes adapt to wind erosion on the active sand dune; and (4) helping psammophytes realize long-distance dispersal.     

The Effects of Habitat on Dispersal Patterns of an Invasive Thistle, Cynara Cardunculus

Dispersal of offspring is a critical step in the spread of invasive species, yet dispersal patterns are seldom well studied, inhibiting effective management and ecological understanding of invasions. Dispersal patterns can be affected by characteristics of the parent plant and by climatic or site characteristics, including community vegetation structure, but these factors have not been studied in the context of plant invasion processes. Cynara cardunculus (L.) is a polycarpic perennial thistle invasive in coastal grasslands in California that produces large numbers of wind-dispersed seeds. This study quantified and compared C. cardunculus dispersal patterns in an exotic grassland (vegetated site) and an agricultural field (non-vegetated site). Seed size variation in C. cardunculus was also quantified within inflorescences, between inflorescences, and between years of production, and compared across dispersal distances. Results indicate that dispersal distance increased dramatically from less than 20 m in the vegetated site to more than 40 m in the non-vegetated site. Plants producing fewer seeds produced heavier seeds and dispersal distance decreased over time, but seed size was not related to dispersal distance, which may serve to spread the risk to seeds or seedlings across the environment. C. cardunculus has great potential for dispersal in open areas such as agricultural fields or disturbed sites, but may be limited in highly structured natural communities. Management of C. cardunculus and other wind-dispersed plant invasions may be improved by prioritizing populations with open or disturbed areas downwind and minimizing the removal of vegetation during dispersal.     

Graphical analysis of the life history evolution of Oenothera glazioviana Micheli

Summary Five-year census data of a semelparous Oenothera glazioviana population in a sand dune system were graphically analyzed. Changes in survivorship until seed production and potential fecundity were calculated from the size-dependent growth, survival and reproduction. The net reproductive rates after logarithmic transformation were plotted against the presumed age of reproduction. The optimal age and size at reproduction that maximize the intrinsic rate of natural increase were determined as a gradient of the tangent drawn from the origin. The natural population reproduced a little earlier than the calculated optimal age but at nearly the optimal size. Examination of the effect of environment on the optimal age and size revealed that an environment imposing higher mortality increases both the optimal age and size, while an environment imposing lower fecundity increases the optimal age but not the optimal size. The relatively large critical size for reproduction of O. glazioviana causes delayed reproduction in infertile environments, which helps the species maintain itself in a risky environment such as the sand dunes.     

A review of the genus Erotendomychus Lea (Coleoptera: Endomychidae)

Species of the Australian genus Erotendomychus are reviewed. The following new species are described and illustrated: E. dentatus, dorrigo, elongatus, emarginatus, erectus, joalah, kirrama, leai, micropunctatus, micrus, ovatus, peckorum, yeatesi. Nomenclatural history, diagnoses and distribution are provided for each species. A key to the species of the genus is presented.     

Factors limiting the distribution ofDigitaria adscendens andEleusine indica on coastal sand dunes in Japan

Fate ofDigitaria adscendens andEleusine indica seedlings under field conditions and their responses to salt spray, drought and nutrients were experimentally investigated in order to evaluate the possible mechanisms controlling the different distributions of the two species in coastal sand dune areas. Salt spray produced no apparent positive effect on the growth or survival of both species. Seeds of each species germinated well in the field, although 80% ofE. indica seedlings died during a summer drought and surviving seedlings neither grew nor bore fruit. The mortality ofD. adscendens seedlings due to the drought was less than 10% and the survivors mostly bore fruit by the end of the growth season. No major difference in the sublethal water saturation deficit was noticed between the two species. However,D. adscendens individuals extended their roots into the deep sandy soil to a much greater extent in water-stressed conditions than in well watered conditions, whereasE. indica showed no such behavior. Additional watering in a dune environment did not help the growth ofE. indica seedlings, but additional nutrients had a markedly stimulatory effect.D. adscendens maintained its growth and fruition with much smaller amounts of nutrients thanE. indica. Soil nitrogen content at a site whereE. indica andD. adscendens were distributed sympatrically was higher than that at a site where onlyD. adscendens was present. Based on these findings, it is proposed thatE. indica seedlings are unable to become established because of their lower resistance to summer drought and the poor nutrient conditions present in a coastal rear sand dune habitat. Dedicated to Prof. emeritus Toshiro Saeki for his fruitful career in plant ecology.     

Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

Reproductive biology and pollen flow of a rheophytic aroid, Furtadoa sumatrensis (Araceae) in the Malesian wet tropics

 A rheophytic aroid, Furtadoa sumatrensis M. Hotta, showed protogynous flowering behaviour, which reduced autogamous pollination within the spadix and promoted outcrossing among spadices by Colocasiomyia sp. (Diptera, Drosophilidae). Self-compatibility of this species allowed geitonogamous crossing between genetically homogeneous spadices from the same individual. Chemical dyes as pollen analogues were carried by Colocasiomyia flies bidirectionally along the forest stream habitat, and their movements were significantly affected by the density of mature spadices in a population. This correlation was consistent among populations. In a population with higher density, the dye's movements were restricted to a smaller distance logarithmically. Estimated neighborhood size (N _e ) and area (A) showed positive and negative correlation with density, respectively. The range of pollen flow was estimated at most within a river system. Received March 7, 2000 Accepted January 23, 2001     

Myrmecochory in Polygala vulgaris L., Luzula campestris (L.) DC. and Viola curtisii Forster in a Dutch dune area

Summary Seed dispersal by ants in Polygala vulgaris, Luzula campestris and Viola curtisii was studied in a primary dune valley on the island of Terschelling, The Netherlands. Normally developed seeds of all three species are taken by the ants into their nests. The ants show a distinct preference for the seeds of the specialized myrmecochore Polygala vulgaris, as compared with the two diplochorous species. It could be demonstrated that the elaiosome is the attractive part of the seed. Mapping studies demonstrate that the dispersal of the seeds by ants has a marked effect on the distribution pattern of the standing population of Polygala and Viola. Adult plants are often found on or close to the active nest mounds of all ant species present, while the growing sites of juvenile individuals and seedlings are practically restricted to the nest environment. The nests of two of the seed-dispersing ant species, viz., those of Lasius niger and Tetramorium caespitum, show differences in soil chemistry with the surroundings. The ant nests are significantly richer in some essential plant macronutrients, such as phosphate, potassium and nitrate. The advantage of myrmecochory in the dune area of Terschelling is discussed.