Performance of reciprocally sown populations of Senecio vulgaris from ruderal and agricultural habitats

Senecio vulgaris from ruderal habitats may colonise crop fields and persist in the new environment. Persistence may occur through phenotypic plasticity or genetic differentiation. In the presence of genetic variation, differential responses of life history traits to selection may lead to local adaptation. A reciprocal seed transplant experiment was conducted to determine environmental and genetic variation of life history traits in S. vulgaris from ruderal and agricultural habitats, and whether infection by the rust Puccinia lagenophorae is a selection factor in S. vulgaris populations. Emergence, growth and fecundity showed environmental as well as genetic variation, as shown by a significant site and origin effect. However, there was no evidence of local adaptation, as indicated by the absence of significant origin by site interactions. Genetic variation in emerging seedling numbers seems to be important for S. vulgaris, an annual plant which has no means other than seed production for propagation. Seedling survival was solely under environmental control stressing the importance of stochastic events for plant mortality. Most S. vulgaris from ruderal sites showed reduced growth, but still reached reproduction, suggesting that S. vulgaris is following the plastic strategy of a stress tolerator to endure unfavourable environments. Plants at most agricultural sites behaved like r-strategists, exploiting a productive environment for rapid plant growth and maximising reproduction. Emergence, survival, growth and fecundity were higher at the agricultural sites. Infection by the rust occurred at all sites, with the highest incidence at the agricultural sites, and was solely determined by the environment. A higher incidence did not result in an increased disease impact on plant growth and reproduction nor did it affect survival of S. vulgaris. Although the majority of life-history traits studied showed phenotypic and genetic variation, which can both be subject to natural selection, no adaptation of S. vulgaris to its local environment was detected.     

Highly Replicated Evolution of Parapatric Ecotypes

Parallel evolution of ecotypes occurs when selection independently drives the evolution of similar traits across similar environments. The multiple origins of ecotypes are often inferred based on a phylogeny that clusters populations according to geographic location and not by the environment they occupy. However, the use of phylogenies to infer parallel evolution in closely related populations is problematic because gene flow and incomplete lineage sorting can uncouple the genetic structure at neutral markers from the colonization history of populations. Here, we demonstrate multiple origins within ecotypes of an Australian wildflower, Senecio lautus. We observed strong genetic structure as well as phylogenetic clustering by geography and show that this is unlikely due to gene flow between parapatric ecotypes, which was surprisingly low. We further confirm this analytically by demonstrating that phylogenetic distortion due to gene flow often requires higher levels of migration than those observed in S. lautus. Our results imply that selection can repeatedly create similar phenotypes despite the perceived homogenizing effects of gene flow.     

Competition, predation, and the distributions of four desert anurans

Several studies have shown that larval competition and susceptibility to predation affect distributions of amphibian assemblages across ephemeral and perennial habitats. However, few studies have examined mechanisms affecting distribution patterns and site use of anurans adapted to highly ephemeral habitats. This study examines hypotheses about competition and predation as mechanisms creating non-overlapping patterns of site use in four anurans that breed in highly ephemeral habitats: Scaphiopus couchii (Couch's spade-foot toad), Gastrophryne olivacea (narrow-mouthed toad), Bufo speciosus (Texas toad), and Bufo punctatus (red-spotted toad). These four anurans showed a significantly nonrandom pattern of co-occurrence. Only 12% of 95 ephemeral breeding sites surveyed were occupied by more than one species. We tested the hypotheses that non-overlapping use of breeding sites was due to activity rates of their tadpoles that in turn reflect their competitive ability and susceptibility to predation. Tadpoles of S. couchii were significantly more active and more susceptible to predation than were tadpoles of G. olivacea, B. speciosus, and B. punctatus. The masses of G. olivacea, B. speciosus, and B. punctatus were less when they were reared with S. couchii, demonstrating the possible competitive dominance of S. couchii. These results suggest that the competitive ability of S. couchii may play a role in excluding G. olivacea, B. speciosus, and B. punctatus from very ephemeral breeding sites, and that susceptibility to predation could play a role in excluding S. couchii from breeding sites of longer duration that are more likely to be colonized by aquatic predators.     

Long-term dynamics of the distribution of the invasive Argentine ant, Linepithema humile, and native ant taxa in northern California

Invasive species, where successful, can devastate native communities. We studied the dynamics of the invasive Argentine ant, Linepithema humile, for 7 years in Jasper Ridge, a biological preserve in northern California. We monitored the distributions at the hectare scale of native ant taxa and L. humile in the spring and fall from 1993 to 1999. We also studied the invasion dynamics at a finer resolution by searching for ants in 1-m² plots. Our results are similar at both scales. The distributions of several native species are not random with regard to L. humile; the distributions of several epigeic species with similar habitat affinities overlap much less frequently than expected with the distribution of L. humile. We found that season had a significant influence on the distributions of L. humile and several native taxa. Over the 7-year period, L. humile has increased its range size in Jasper Ridge largely at the expense of native taxa, but there is seasonal and yearly variation in this rate of increase. Studies of invasions in progress which sample across seasons and years may help to predict the spread and effects of invasive species.     

Molecular evidence that fireweed (Senecio madagascariensis, Asteraceae) is of South African origin

ITS1 sequence data was obtained for fireweed (Senecio madagascariensis) andS. lautus from Australia,S. madagascariensis andS. inaequidens from South Africa andS. madagascariensis from Madagascar. Despite the low level of variation (0.0–3.4%), these data further resolve the controversy concerning the identity and origin of fireweed. They confirm that fireweed is part of the South AfricanS. madagascariensis/S. inaequidens complex, and indicate that the infestation in Australia originated from South Africa as opposed to Madagascar. This will facilitate a resumption of biological control efforts in Australia and will direct surveys for control agents to South Africa.     

Role of mucilage in the germination of Artemisia sphaerocephala (Asteraceae) achenes exposed to osmotic stress and salinity

Artemisia sphaerocephala (Asteraceae) is one of the pioneer species in moving and semi-stable sand dunes in the deserts of northwest China. The outer surface of A. sphaerocephala achenes contains a pectinaceous mucilage layer that can imbibe a large amount of water when wetted. We hypothesized that the mucilage can aid achene germination in heterogeneous environments. Germination of both intact achenes and those from which the mucilage had been removed (demucilaged) declined with increasing osmotic potential and NaCl concentration. However, the germination percentage of intact achenes was significantly higher than that of demucilaged achenes. The early seedling growth of intact achenes did not differ significantly from that of demucilaged achenes in either osmotic potential or NaCl solutions. Achene mucilage presumably plays an ecologically important role in the life cycle of A. sphaerocephala by aiding germination in osmotically- and saline-stressful habitats of the cold desert environment.     

Dispersion,germination and growth of seedlings ofSagittaria lancifolia L.

Sagittaria lancifolia L. a perennial helophyte growing in shallow waters has asexual reproduction by rhizomes and sexual reproduction by achenes. We studied the morphological characteristics of achenes related to its dispersion, some factors affecting its germination and the development stages and growth of seedlings. This species produces many achenes, and disperses through epizoochory and hydrochory. Achenes germinate on light only. 100% germination was obtained at 25°C. Under conditions of inundation the germination period decreases, and the percentage of germination in creases when compared to achenes with or without imbibition. Achenes in anaerobic conditions have reduced germination. The growth of seedlings ofS. lancifolia was affected by the temperature: each organ of the seedlings grows differently in dependence upon this factor. For example, roots grow faster during the early stages of the seedlings.     

Molluscan assemblages on coral reefs and associated hard substrata in the northern Red Sea

Information on spatial variability and distribution patterns of organisms in coral reef environments is necessary to evaluate the increasing anthropogenic disturbance of marine environments (Richmond 1993; Wilkinson 1993; Dayton 1994). Therefore different types of subtidal, reef-associated hard substrata (reef flats, reef slopes, coral carpets, coral patches, rock grounds), each with different coral associations, were investigated to determine the distribution pattern of molluscs and their life habits (feeding strategies and substrate relations). The molluscs were strongly dominated by taxa with distinct relations to corals, and five assemblages were differentiated. The Dendropoma maxima assemblage on reef flats is a discrete entity, strongly dominated by this encrusting and suspension-feeding gastropod. All other assemblages are arranged along a substrate gradient of changing coral associations and potential molluscan habitats. The Coralliophila neritoidea-Barbatia foliata assemblage depends on the presence of Porites and shows a dominance of gastropods feeding on corals and of bivalves associated with living corals. The Chamoidea-Cerithium spp. assemblage on rock grounds is strongly dominated by encrusting bivalves. The Drupella cornus-Pteriidae assemblage occurs on Millepora-Acropora reef slopes and is strongly dominated by bivalves associated with living corals. The Barbatia setigera-Ctenoides annulata assemblage includes a broad variety of taxa, molluscan life habits and bottom types, but occurs mainly on faviid carpets and is transitional among the other three assemblages. A predicted degradation of coral coverage to rock bottoms due to increasing eutrophication and physical damage in the study area (Riegl and Piller 2000) will result in a loss of coral-associated molluscs in favor of bivalve crevice dwellers in dead coral heads and of encrusters on dead hard substrata.     

Natural selection drives leaf divergence in experimental populations of Senecio lautus under natural conditions

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Variability and Cryptic Heteromorphism of Ambrosia artemisiifolia Seeds: What Consequences for its Invasion in France?

BACKGROUND AND AIMS: Ambrosia artemisiifolia is a ruderal weed introduced from North America to Europe. It produces large amount of achenes which are highly heterogeneous in size. Due to the preponderant role of propagules in invasive plant processes, the achene mass variability related to germination, dispersal strategy and life history traits of offspring were investigated within this species. METHODS: The variability in achene mass was quantified among six populations sampled in different habitats. The effects of achene mass variation on germination were studied. The percentages of floating and non-floating achenes were evaluated in the studied populations. The consequences of floatability on the growth and traits of the offspring were studied. KEY RESULTS: Mean achene mass ranged from 1.72 to 3.60 mg, depending on the populations, and was highly variable. Variation among achenes within plants accounted for 63.9 % of the variance, whereas variances among plants within each population (22.2 %) and among populations (13.9 %) were lower. Achene masses were also positively correlated to the total germination percentage for four populations out of six. Two kinds of achenes were distinguished: floating and non-floating. The majority of floating achenes (90 %) sank 24 h after water immersion. Whatever the population, floating achenes were lighter, more dormant and germinated faster than non-floating achenes. Plants which issued from floating achenes had better growth than those from non-floating achenes. CONCLUSIONS: The capacity of A. artemisiifolia to be invasive in Europe appears to be high, possibly due to its huge plasticity in seed mass which may help it to cope with a wide range of conditions and to establish in disturbed habitats. Furthermore, the recent invasion of southern France by A. artemisiifolia could be partially explained by water dispersal of achenes through rivers and has pinpointed its colonization potential along French rivers.     

Photosynthesis, photoinhibition, and nitrogen use efficiency in native and invasive tree ferns in Hawaii

Photosynthetic gas exchange, chlorophyll fluorescence, nitrogen use efficiency, and related leaf traits of native Hawaiian tree ferns in the genus Cibotium were compared with those of the invasive Australian tree fern Sphaeropteris cooperi in an attempt to explain the higher growth rates of S. cooperi in Hawaii. Comparisons were made between mature sporophytes growing in the sun (gap or forest edge) and in shady understories at four sites at three different elevations. The invasive tree fern had 12-13 cm greater height increase per year and approximately 5 times larger total leaf surface area per plant compared to the native tree ferns. The maximum rates of photosynthesis of S. cooperi in the sun and shade were significantly higher than those of the native Cibotium spp (for example, 11.2 and 7.1 µmol m^-2 s^-1, and 5.8 and 3.6 µmol m^-2 s^-1 respectively for the invasive and natives at low elevation). The instantaneous photosynthetic nitrogen use efficiency of the invasive tree fern was significantly higher than that of the native tree ferns, but when integrated over the life span of the frond the differences were not significant. The fronds of the invasive tree fern species had a significantly shorter life span than the native tree ferns (approximately 6 months and 12 months, respectively), and significantly higher nitrogen content per unit leaf mass. The native tree ferns growing in both sun and shade exhibited greater photoinhibition than the invasive tree fern after being experimentally subjected to high light levels. The native tree ferns recovered only 78% of their dark-acclimated quantum yield (F_v/F_m), while the invasive tree fern recovered 90% and 86% of its dark-acclimated F_v/F_m when growing in sun and shade, respectively. Overall, the invasive tree fern appears to be more efficient at capturing and utilizing light than the native Cibotium species, particularly in high-light environments such as those associated with high levels of disturbance.     

Predation environment predicts divergent life-history phenotypes among populations of the livebearing fish Brachyrhaphis rhabdophora

We document a strong association between predation environment and life-history phenotypes in the Costa Rican livebearing fish Brachyrhaphis rhabdophora. Populations that co-occurred with piscine predators attained maturity at a smaller size, and produced more, smaller offspring relative to populations from predator-free environments. These differences persisted over 3 years and between wet and dry seasons within a year. Reproductive allotment did not differ between predation environments, but was greater in the wet season than in the dry season. We also examined the phenotypic covariance structure among life-history traits and found traits to be highly correlated. Based on life-history differences, discriminant analyses showed that populations could be neatly classified by predation category, and could be reasonably classified into wet and dry season categories. Finally, we found that the pattern of predator-associated life-history divergence in B. rhabdophora is remarkably similar to that of the taxonomically distinct Trinidadian guppy (Poecilia reticulata), possibly pointing to an evolutionary convergence between these two systems.     

The offspring size/fecundity trade-off and female fitness in the Atlantic molly (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae)

Across a variety of taxa, large offspring have been demonstrated to have a fitness advantage over smaller offspring of the same species. However, producing large offspring often comes at the cost of having to produce fewer young, and the payoff (and thus, evolutionary outcome) of this trade-off is expected to vary between environments. Atlantic mollies (Poecilia mexicana: Poeciliidae, Teleostei), inhabiting a sulfidic cave and various non-sulfidic surface habitats in Tabasco (Mexico), are reproductively isolated and evolved divergent female life-history traits: females of the cave ecotype produce considerably fewer, but larger offspring. Stressful (sulfidic) environments may favor the production of larger offspring, as they are better able to cope with chemical stressors. It remains to be determined though if increased offspring survival outweighs the fitness cost of producing fewer but larger offspring even under benign laboratory conditions. We tested 30-day newborn survival of offspring from wild-caught P. mexicana females from diverging populations in a low-density, no predation, no cannibalism, and ad-libitum-food, benign laboratory environment. Survival rates were highly skewed towards larger cave molly offspring; however, surface molly females still had a higher fitness than cave molly females in terms of higher total numbers of surviving offspring. Our study provides evidence for an innate fitness advantage of larger cave molly offspring. Furthermore, the observed differences in life-history strategies could promote further divergence and reproductive isolation among these ecotypes of P. mexicana, because cave molly females immigrating into the adjacent surface habitats would most likely be selected against.     

Biological characteristics of an invasive south African species

Senecio inaequidens, a south African species that has recently invaded parts of Europe, especially human disturbed habitats, was examined under laboratory and glasshouse conditions to assess the germination of achenes, soil seed banks, growth of seedlings and its breeding system. These observations allowed identification of ecological characteristics that may contribute to the invasiveness of the species. Achenes derived from three populations of S. inaequidens were germinated over a wide temperature range (from 14/6 °C day/night temperatures to a constant temperature of 30 °C). They exhibited a rapid achene germination especially when temperatures reached 20 °C. Germination of seeds from soil samples also exhibited a prolonged germination over time suggesting therefore different types of dormancy. Growth response was studied along a nitrogen gradient from 0.33 to 16.0 mmol l^–1 and data were interpreted using growth analysis. Seedlings of S. inaequidens exhibited a slow growth in low nutrient availability and a rapid growth rate when exposed to high nutrient availability, which contributed to producing a large leaf area and biomass. Four types of pollination were studied in S. inaequidens. Hand-pollinations showed that S. inaequidens was mostly self-incompatible. Self-pollinated capitula only showed 7.6% viable achenes, significantly lower than insect-pollinated and cross-pollinated capitula. Some individuals were more self-fertile than others ensuring the production of some offspring in a situation of colonisation. Natural pollination occurred with generalist insects. Most of these results confirm the invasive ability of S. inaequidens.     

Dormancy and dispersal in dimorphic achenes of tansy ragwort,Senecio jacobaea L. (Compositae)

Summary Marginal and central florets of the capitula of tansy ragwort Senecio jacobaea yield different kinds of fruit. The central (disk) achenes are lighter (x±SE=199±5g), more numerous (x±SE=58±0.6 achenes per head), and are equipped with a pappus aiding wind transport and rows of trichomes aiding animal transport. The marginal (ray) achenes are heavier (x±SE=286±7g), less numerous (virtually invariant at 13 achenes per head), and lack dispersal structures. Whereas disk achenes are relased shortly after they mature, ray achenes are retained by the parent for a period of months following maturity.Germination at constant temperature (20°C) and with alternating light (12 h light: 12 h dark) demonstrated that disk and ray achenes exhibit different germination syndromes. Germination percentage increases linearly with achene fresh weight in both types; for a given weight, disk achenes have a higher germination percentage than ray achenes. Germination time decreases with increasing achene weight in disk achenes, but increases with achene weight in ray achenes.The germination percentages and germination times for disk and ray achenes diverge progressively with increasing achene weight. The divergence in behavior is a result of diverging patterns of dry matter allocation in the two achene types. Increase in the size of disk achenes favors the embryo fraction, thereby speeding germination while reducing protection. Increase in the size of the ray achenes favors the pericarp fraction, thereby increasing protection while delaying germination.Reduced germination percentage and germination speed of the ray achenes were shown by experimental manipulation to be caused by physical inhibition by their thicker pericarps.Dimorphism in ragwort likely speards germination out in space and time, thereby increasing the number of safe sites an individual parent can exploit in disseminating offspring. The syndrome in other heteromorphic composites resembles that of ragwort, generally combining reduced dispersal-delayed germination in the outer achenes and distance dispersal-quick germination in the central achenes. The outer achenes are generally less numerous and larger. Dispersal traits (large numbers, early release and light wieght) are the direct opposite of dormancy traits (small numbers, delayed release and heavier weight). Thus conflicts between the properties determining dormancy and dispersal appear to require separate dormancy and dispersal phenotypes.     

Plant diversity,herbivory and resistance of a plant community to invasion in Mediterranean annual communities

Several components of the diversity of plant communities, such as species richness, species composition, number of functional groups and functional composition, have been shown to directly affect the performance of exotic species. Exotics can also be affected by herbivores of the native plant community. However, these two possible mechanisms limiting invasion have never been investigated together. The aim of this study was to investigate the relationships between plant diversity, herbivory and performance of two annual exotics, Conyza bonariensis and C. canadensis, in Mediterranean annual communities. We wanted to test whether herbivory of these exotics was influenced either by species richness, functional-group richness or functional-group composition. We also studied the relationship between herbivory on the exotic species and their performance. Herbivory increased with increasing species and functional-group richness for both Conyza species. These patterns are interpreted as reflecting a greater number of available herbivore niches in a richer, more complex, plant community. The identities of functional groups also affected Conyza herbivory, which decreased in the presence of Asteraceae or Fabaceae and increased in the presence of Poaceae. Increasing herbivory had consequences for vegetative and demographic parameters of both invasive species: survival, final biomass and net fecundity decreased with increasing herbivory, leading to a loss of reproductive capacity. We conclude that communities characterised by a high number of grass species instead of Asteraceae or Fabaceae may be more resistant to invasion by the two Conyza species, in part due to predation by native herbivores.     

Secondary invasion of <Emphasis Type="Italic">Acer negundo</Emphasis>: the role of phenotypic responses versus local adaptation

During plant species invasions, the role of adaptive processes is particularly of interest in later stages of range expansion when populations start invading habitats that initially have not been disposed to invasions. The dioecious tree Acer negundo, primarily invasive in Europe in wet habitats along riversides and in floodplains, has increased its abundance in dry habitats of industrial wasteland and ruderal sites during the last decades in Eastern Germany. We chose 21 invasive populations from wet and from dry habitats in the region of Halle, Saxony-Anhalt, Germany, to test whether Acer negundo exhibits a shift in life-history strategy during expansion into more stressful habitats. We analyzed variables of habitat quality (pH, soil moisture, exchangeable cations, total C and N content) and determined density, sex ratio and regeneration of the populations. In addition, we conducted germination experiments and greenhouse studies with seedlings in four different soil moisture environments. Local adaptation was studied in a reciprocal transplant experiment. We found habitat type differentiation with lower nutrient and water supply at the dry sites than at the moist sites and significant differences in the number of seedlings in the field. In accordance, seeds from moist habitats responded significantly faster to germination treatments. In the transplant experiment, leaf life span was significantly larger for populations originating from dry habitat types than from moist habitats. This observed shift in life history strategy during secondary invasion of A. negundo from traits of establishment and rapid growth towards traits connected with persistence might be counteracted by high gene flow among populations of the different habitat types. However, prolonged leaf life span at dry sites contributed remarkably to the invasion of less favourable habitats, and, thus, is a first indication of ongoing adaptation.     

Overcoming seed dormancy in ex situ plant germplasm conservation programmes; an example in the endemic Argyranthemum (Asteraceae: Anthemideae) species from the Canary Islands

Germplasm of 21 diverse Argyranthemum taxa was collected from contrasting ecological zones in the Canary Islands. Seed dormancy was considerable in the majority of taxa. Extensive investigations, based on a germination test procedure algorithm for Asteraceae, with achenes from ray and disc florets of five contrasting taxa identified a procedure to promote full (85%) germination of the seeds from both ray and disc florets of all five taxa; viz, excision of the seeds from the achenes, followed by testing at 15°C with 2.6×10^-3 m GA₃ co-applied. Subsequent tests showed that this regime was effective in promoting full germination in seeds from both ray and disc florets of the remaining 16 taxa. The results are discussed in the context of ex situ plant germplasm conservation.     

Tetraploid European Salicornia species are best interpreted as ecotypes of multiple origin

Salicornia procumbens and S. stricta are two tetraploid European salt marsh species of locally adjacent but ecologically differentiated distribution. Whereas S. procumbens grows in the lowest part of the salt marsh, it is replaced by S. stricta in the middle part (and diploid Salicornias in the upper part). Using AFLPs and a reciprocal transplantation experiment, we investigated whether the two species represent distinct evolutionary lineages. The analysis of AFLP variation clearly showed that both species are not monophyletic. Also, accessions do not cluster according to geographical origin. The transplantation experiment revealed that S. procumbens shows significantly reduced fitness when transplanted into the habitat of S. stricta. Reduction of fitness of S. stricta in the habitat of S. procumbens is less obvious. In both habitats S. procumbens performs better than S. stricta in terms of seed production and dry weight. On the background of these results, it is hypothesized that S. procumbens and S. stricta represent intraspecific ecotypes which originated repeatedly in adaptation to their specific environments and which dispersed widely. We hypothesize that the ecologically vicariant distribution of S. procumbens and S. stricta along the inundation gradient in the saltmarsh in spite of their good performance in the respective non-native habitats results from strong selection during seed germination and seedling establishment. Reproductive isolation between locally adjacent or even intermingled ecotypes is suggested to be achieved by high levels of selfing.     

The ecology of germination and reproduction of less frequent and vanishing species of the Czechoslovak flora II.Pulsatilla slavica Reuss

The conditions of embryo development, of germination and sprouting in the speciesPulsatilla slavica (north Carpathian endemite) were ascertained. By studies of germination ecology ofPulsatilla slavica it was found that the development of the underdeveloped embryo and germination set in at about 10 °C to about 30 °C. Of the temperatures examined, the temperature of about 25 °C, and alternating ones, the temperature of 10/30 °C appeared to be the optima. At the temperature of 25 °C, the rate of loss of germination capacity was ascertained partly in fully ripe achenes, partly in achenes collected in the stage of wax-ripeness. After 39 weeks of room storage, 62.6% of fully ripe achenes and 32.6% of achenes collected in the stage of waxripeness germinated. The achenes sown after full ripeness in the middle of June sprouted in the middle of July, those sown at the beginning of October sprouted during the first decade of May, and those sown at the beginning of April sprouted during the, third decade of May-Pulsatilla slavica reproduces spontaneously only in the generative way, but it can also be propagated vegetatively by root cuttings.