Ploidy level and origin of the European invasive weed Senecio inaequidens (Asteraceae)

Native to South-Africa, species of the Senecio inaequidens complex are presently invasive in Europe, Australia and South-America. Previously, different ploidy levels have been found in these different areas, with only tetraploid individuals reported in Europe, and only diploids in South-Africa and Australia. In the present study chromosome counts and flow cytometry were used to survey DNA ploidy levels in a large sample of 66 native and 21 European invasive populations. One Mexican individual was also added to the study. We found only tetraploid individuals occurring in Europe, whereas both ploidy levels, diploid and tetraploid, were found in South-Africa. Moreover, based on genome size, we suggest that two largely allopatric varieties of diploids exist in South-Africa. The Mexican individual was diploid. We suggest that European tetraploid individuals come from South-Africa and hypothesize that a hybridization event between the two DNA types of diploids occurred in the Lesotho area. The taxonomic difficulties surrounding species of theS. inaequidens complex are briefly discussed.     

Increase in mate availability without loss of self-incompatibility in the invasive species Senecio inaequidens (Asteraceae)

The evolution of the strength of self-incompatibility in Senecio inaequidens , a native of South Africa was investigated in relation to its invasion in Europe. Levels of self-incompatibility were estimated with hand-pollinations in five populations in greenhouse conditions. One population came from the native range of the species and four populations were sampled in Europe from two independent transects of colonization with old and recent populations. Contrary to Baker's law predictions, our results suggest that the species has a sporophytic self-incompatible system maintained in all populations. We suggest that the ability of S. inaequidens to colonize new sites with a self-incompatibility system is promoted by its ecological characteristics (perenniality, extended reproductive period, massive seed production, generalist pollinators). In addition, we found that mate availability was increased (1) in the introduced range compared to the native range, (2) in marginal versus central European populations. Possible explanations for this surprising result are discussed.     

The effects of herbivory and competition on the invasive alien plant Senecio inaequidens (Asteraceae)

Abstract. Senecio inaequidens DC. (Asteraceae) is an invasive alien plant introduced to Europe from South Africa in around 1896. It contains pyrrolizidine alkaloids that are toxic to livestock and humans. S. inaequidens would therefore be an economic and ecological problem if it became established and abundant in natural or farmed grassland ecosystems. We conducted field experiments using a split‐plot design to determine the effects of rabbit grazing, interspecific plant competition, mollusc and insect herbivory on growth, survival and reproduction of S. inaequidens. Plants were grown from seeds of three different ecotypes under standardized greenhouse conditions and transplanted into field plots. Rabbits (Oryctolagus cuniculus L.) were excluded from experimental plots using rabbit fences. Competition was manipulated by either creating subplots with bare ground or leaving the vegetation cover intact. Data were recorded between June and August 2002. Ecotypes differed significantly in morphological parameters, and in their responses to invertebrate herbivory. Interspecific plant competition and rabbit grazing significantly reduced growth and reproduction of S. inaequidens. Regrowth shoots of S. inaequidens produced after rabbit grazing were not subsequently eaten by rabbits. Unpalatability of regrowth shoots may be attributable to changes in pyrrolizidine alkaloid composition with plant age. Mollusc herbivory significantly reduced the number of capitulae produced. We found adults of Longitarsus jacobaeae Waterhouse (Coleoptera: Chrysomelidae), a specialist herbivore of European Senecio jacobaea L. (Asteraceae), feeding on 79% of S. inaequidens plants. 320 larvae of Tyria jacobaeae L. (Lepidoptera: Arctiidae) did not feed on S. inaequidens under free‐choice field conditions. We conclude that S. inaequidens is able to survive and reproduce in disturbed grassland ecosystems. L. jacobaeae might be a suitable agent for biological control of S. inaequidens in European introduced populations in the future.     

Seasonal variability in pyrrolizidine alkaloids in Senecio inaequidens from the Val Venosta (Northern Italy)

Senecio inaequidens is a neophyte originating from South Africa that has managed to spread to Europe and colonize large areas. This plant has toxic pyrrolizidine alkaloids (PAs) that represent a health risk to humans and animals. The aim of this work was to measure the content of PAs of these plants in continuous intervals over the entire growing season. The plants were separated into their organs, such as sprouts, stems, leaves and inflorescences. PAs were extracted from the dried plants using an acidic methanol solution, purified and measured by GC/MS. The average PA content of plants over the growing season was 0.33% of dry weight. The highest PA levels were about 1% in the dry weight and were found in the young sprouts and flower heads. There were nine PAs present, of which six could be identified. The main alkaloid was retrorsine followed by senecivernine, senecionine, integerrimine, usaramine and seneciphylline. Due to the high PA content of the inflorescences, the long flowering period and the rapid invasion dynamics, this species presents a high health risk for humans and animals.     

Evolution of dispersal traits along an invasion route in the wind‐dispersed Senecio inaequidens (Asteraceae)

In introduced organisms, dispersal propensity is expected to increase during range expansion. This prediction is based on the assumption that phenotypic plasticity is low compared to genetic diversity, and an increase in dispersal can be counteracted by the Allee effect. Empirical evidence in support of these hypotheses is however lacking. The present study tested for evidence of differentiation in dispersal‐related traits and the Allee effect in the wind‐dispersed invasive Senecio inaequidens (Asteraceae). We collected capitula from individuals in ten field populations, along an invasion route including the original introduction site in southern France. In addition, we conducted a common garden experiment from field‐collected seeds and obtained capitula from individuals representing the same ten field populations. We analysed phenotypic variation in dispersal traits between field and common garden environments as a function of the distance between populations and the introduction site. Our results revealed low levels of phenotypic differentiation among populations. However, significant clinal variation in dispersal traits was demonstrated in common garden plants representing the invasion route. In field populations, similar trends in dispersal‐related traits and evidence of an Allee effect were not detected. In part, our results supported expectations of increased dispersal capacity with range expansion, and emphasized the contribution of phenotypic plasticity under natural conditions.     

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.     

The control of chiasma frequeney within a polyploid series in the genus Senecio (Compositae)

Chiasma frequency in a polyploid series, and in the hybrids formed within it, are shown to be directly related to ploidy level, and to be independent of the apparent effective synapsis. The observations are related to previous observations and models on the control of chiasma frequency.     

A novel method to infer the origin of polyploids from Amplified Fragment Length Polymorphism data reveals that the alpine polyploid complex of Senecio carniolicus (Asteraceae) evolved mainly via autopolyploidy

Despite its evolutionary and ecological relevance, the mode of polyploid origin has been notoriously difficult to be reconstructed from molecular data. Here, we present a method to identify the putative parents of polyploids and thus to infer the mode of their origin (auto‐ vs. allopolyploidy) from Amplified Fragment Length Polymorphism (AFLP) data. To this end, we use Cohen's d of distances between in silico polyploids, generated within a priori defined scenarios of origin from a priori delimited putative parental entities (e.g. taxa, genetic lineages), and natural polyploids. Simulations show that the discriminatory power of the proposed method increases mainly with increasing divergence between the lower‐ploid putative ancestors and less so with increasing delay of polyploidization relative to the time of divergence. We apply the new method to the Senecio carniolicus aggregate, distributed in the European Alps and comprising two diploid, one tetraploid and one hexaploid species. In the eastern part of its distribution, the S. carniolicus aggregate was inferred to comprise an autopolyploid series, whereas for western populations of the tetraploid species, an allopolyploid origin involving the two diploid species was the most likely scenario. Although this suggests that the tetraploid species has two independent origins, other evidence (ribotype distribution, morphology) is consistent with the hypothesis of an autopolyploid origin with subsequent introgression by the second diploid species. Altogether, identifying the best among alternative scenarios using Cohen's d can be straightforward, but particular scenarios, such as allopolyploid origin vs. autopolyploid origin with subsequent introgression, remain difficult to be distinguished.     

The Influence of Leaf "Windows" on Crassulacean Acid Metabolism in the South African Succulent Senecio Rowleyanus (Asteraceae)

Net CO₂ exchange rate (P_N) of shoots and diel fluctuations in titratable acidity of leaves of Senecio rowleyanus were measured to determine whether penetration of radiant energy through leaf “windows” (narrow, translucent strips on the leaf epidermis) resulted in increased CAM. Nocturnal P_N and nighttime increases in acidity were compared among plants with windows covered with reflective adhesive tape, transparent adhesive tape (to control for potential effects of the adhesive), and no tape. The windows did not significantly enhance the degree of CAM in S. rowleyanus. This revised version was published online in September 2006 with corrections to the Cover Date.     

Monitoring and modeling the invasion of the fast spreading alien Senecio inaequidens DC. in an alpine region

We modeled the distribution of the South African alien Senecio inaequidens DC. in the Aosta Valley, Western Italian Alps, using data extracted from the Regional floristic database and from an intensive field survey carried out in years 2009–2010. The aims of the work were (1) to evaluate whether the species is in the introduction, colonization, or establishment stage of invasion, (2) to detect the environmental factors that drive the invasion process, and (3) to highlight the potential range of distribution of the alien species. The modeling framework was a stepwise generalized linear model (GLM), using gridded presence/absence data and environmental predictors such as topography, climate, land use, and anthropogenic and natural disturbances. GLM were fit both with and without an additional independent variable to take into account current dispersal limitations. S. inaequidens displayed a very fast spread in the Aosta Valley in the years 1990–2010. The species was positively associated with roads and rivers, southern slopes, and negatively with elevation. However, it was found at an elevation of 1600 m, showing the ability to reach higher elevations than those observed for other invasive alien species, and confirming to be pre-adapted to mountain conditions. The difference between the species distribution models, with and without dispersal constraints, suggested that the availability of seed sources still limits the potential distribution of the species, rather than the environmental variables, and that the realized regional niche differs to a great extent from the equilibrium niche. When limitations to the seed source cease (i.e., in the establishment stage), the species will likely invade large areas that are currently characterized by pastures and grasslands with native species of high agricultural importance. The invasion of S. inaequidens should therefore be considered a serious threat, due to its potential to invade mountain regions, and in particular to colonize habitats used for grazing and forage, thus leading to a high risk for cattle and human health. We discuss the relevance of the results both concerning communication with the public and to support local eradication and control activities. The inclusion of S. inaequidens in the “black list” of the regional law for the conservation of alpine flora (L.R. 45/2009) will help to transfer the information and support invasion control, in particular at medium elevations.     

Factors affecting the invasion success of Senecio inaequidens and S. pterophorus in Mediterranean plant communities

Question: Plant invasions result from complex interactions between species traits, community characteristics and environmental variations. We examined the effect of these interactions on the invasion potential of two invasive Senecio species, S. inaequidens and S. pterophorus, across three Mediterranean plant communities in a natural park. Location: Catalonia, NE Spain. Methods: We carried out two series of experimental seedling transplantations, in the spring and fall of 2003, in grassland, shrubland and Quercus ilex forest. Competition with neighbouring plants and water availability were manipulated. We evaluated the survival, growth and reproduction with respect to each treatment combination. Results: Any habitat can be colonised if disturbance occurs. In the absence of disturbance, shrubland enhanced the survival of seedlings. Competition with resident vegetation dramatically reduced survival in grassland and forest when establishment occurred in the spring. However, establishment in the fall promoted invasion in grassland and shrubland, even in the undisturbed treatment. Grassland allowed the highest growth and reproductive performance of both species while forest was the most resistant habitat to invasion. S. inaequidens had a higher growth rate and a shorter pre‐reproductive period than S. pterophorus. S. pterophorus produced more biomass and was more dependent on water availability than S. inaequidens. Conclusions: In the light of our results, we recommend surveying open shrublands and grasslands after periods of rainfall. Special attention should be paid to S. pterophorus, which is currently spreading. A preliminary assessment of the invasive‐ness of this plant is given in this study.     

Differentiation of reproductive and competitive ability in the invaded range of Senecio inaequidens: the role of genetic Allee effects, adaptive and nonadaptive evolution

Genetic differentiation in the competitive and reproductive ability of invading populations can result from genetic Allee effects or r/K selection at the local or range-wide scale. However, the neutral relatedness of populations may either mask or falsely suggest adaptation and genetic Allee effects. In a common-garden experiment, we investigated the competitive and reproductive ability of invasive Senecio inaequidens populations that vary in neutral genetic diversity, population age and field vegetation cover. To account for population relatedness, we analysed the experimental results with 'animal models' adopted from quantitative genetics. Consistent with adaptive r/K differentiation at local scales, we found that genotypes from low-competition environments invest more in reproduction and are more sensitive to competition. By contrast, apparent effects of large-scale r/K differentiation and apparent genetic Allee effects can largely be explained by neutral population relatedness. Invading populations should not be treated as homogeneous groups, as they may adapt quickly to small-scale environmental variation in the invaded range. Furthermore, neutral population differentiation may strongly influence invasion dynamics and should be accounted for in analyses of common-garden experiments.     

Multivariate analysis of morphological and anatomical characters of Calophyllum (Calophyllaceae) in South America

Calophyllum (Calophyllaceae), previously placed in Clusiaceae, is easily recognizable by its opposite entire leaves with close parallel venation alternating with resin canals. However, distinction between species has been difficult, because of infraspecific variation in tepal and stamen number and resemblance among species that share similar habitats. Here, I report the results of multivariate analyses of morphological and anatomical characters for Calophyllum in South America, and provide a taxonomic treatment for the genus in South America, the first since that of Vesque in 1893. Thirteen preliminary morphogroups were identified. Thirty‐two morphological characters of the leaf, flower and fruit from 401 specimens, and 17 anatomical leaf characters from 45 specimens were measured and analysed using principal component analyses (PCAs) and discriminant analyses (DAs). PCAs were used to find groups and DAs were used to validate those PCAs that were potential groups. Two main subgroups were identified in the general analysis. Subgroup M1 has terete stems and smaller leaves and flowers than subgroup M2, which, instead, has quadrangular stems. Only subgroup M2 showed distinctive clusters in regional and local analyses. Distinctive clusters and morphological and anatomical characters helped us to recognize four species in South America, including a new species, Calophyllum pubescens sp. nov. . In addition, a new species, Calophyllum mesoamericanum sp. nov. , is described from Central America. © 2013 The Linnean Society of London,     

Potential spread of the invasive plant Hydrilla verticillata in South Africa based on anthropogenic spread and climate suitability

Hydrilla verticillata (L.f.) Royle is a submerged aquatic plant native to Asia and Australia that is highly invasive in the USA and was first recorded in South Africa in 2006. It is only known from one locality, Pongolapoort Dam in KwaZulu-Natal Province, but there are fears that it might spread to other sites. The primary vector of spread in the USA is recreational boaters and anglers. A survey at a fishing competition on Pongolapoort Dam showed that anglers travel considerable distances around South Africa (73% of water bodies were >200 km, visited by 68% of the respondents). A Threat Index for freshwater bodies throughout South Africa visited by participants of the competition was calculated showing that dams in the vicinity of the infestation were more at risk from invasion. Further, the potential distribution of the weed based on climatic matching with the region of origin showed that most of the country was suitable for establishment, with the exception of the high-lying interior of the country. Recommendations for reducing the potential spread of hydrilla in South Africa are presented.     

Simulated herbivory counteracts the effects of grass competition on the invasive fireweed (Senecio madagascariensis)

Competition, herbivory and their interaction play a significant role in determining the competitive ability and survival of individual plant species. Understanding these processes and interactions can improve the efficacy of biocontrol programs against invasive weeds. Senecio madagascariensis (fireweed) is an invasive weed of South African origin that reduces pastoral productivity and poisons livestock in several countries, notably Australia. Although competitive pastures can suppress the weed’s growth in Australia, its competitive nature is poorly understood in relation to its invasion success. This greenhouse study assessed the growth and reproductive yield of fireweed growing in competition with six native and introduced grasses present in both South Africa and Australia. Since fireweed is a target for biocontrol in Australia, we examined whether its response to grass competition changed with herbivory (simulated by 40% leaf removal). The effect of grass competition and herbivory on the weed’s biomass and floral productivity was examined during a 12‐week pot trial in South Africa. Floral numbers were unaffected by both grass competition and herbivory. Biomass was used to calculate Relative Interaction Indices (RII) to quantify the weed’s competitive or facilitative response. This index compares a specific measurable trait, such as biomass, of fireweed growing alone, to fireweed growing with grass to determine the level of competitive suppression or facilitation resulting from the interaction. Despite the lack of species‐specific effects of grass competition, the presence of grass suppressed fireweed’s foliar, root and whole plant biomass the most when herbivory was absent. With herbivory, fireweed did not suffer from any measurable competitive suppression. This lack of competitive suppression may be due to an induced allelopathic response, given the levels of pyrrolizidine alkaloids common in many Senecio species. Since this result may weaken the case for biocontrol, the weed’s competitive responses should be verified in relation to actual insect herbivory.     

Modeling the spread of invasive nutrias (Myocastor coypus) over Iran

Nutria (Myocastor coypus) is a native aquatic rodent to South America, and was introduced to Europe, Asia, Africa and North America for fur farming. The South American nutria or coypu is now considered a pest in the area of introduction, because of its negative impact on biological diversity and ecological relationships. Having information on the invasion range of exotic species is crucial for understanding the ecology of invasive spread and for making good conservation and management planning to address this problem. At the beginning of the 20th century, nutria was introduced into Asia. Nutria was recorded for the first time in Iran in 1995. In the present study we proposed a multiple spatial scale approach to predict the invasion trends of the nutria in Iran, and to define up the “suitable scale” for predicting the invasion trends of this species. Our results highlighted the importance of environmental variables including vegetation density (for food and nesting) and water resource (streams, rivers, and lakes) in distribution of the nutria. Potential areas for the presence of the nutria are located near the Caspian Sea, west and central Iran which receive more precipitation than other parts of the country. Therefore, these parts of Iran may face a much greater risk of invasion risk in the future. Moreover, these results can show the possible risk of nutria invasion to the northern and western neighbors of Iran.     

Unexpected morphological and karyological changes in invasive Carpobrotus (Aizoaceae) in Provence (S-E France) compared to native South African species

Hybridization processes can lead to evolutionary changes, particularly in co-introduced congeneric plant species, such as Carpobrotus spp. which are recognized as invasive in Mediterranean climate regions. Morphological and karyological comparisons have therefore been made between native Carpobrotus edulis and C. acinaciformis in South Africa and their invasive counterparts in Provence (C. edulis and C. aff. acinaciformis). Morphological data exhibited the most significant differences in invasive C. aff. acinaciformis that forms a new phenotypic variant. Unexpected chromosomal restructuring has been highlighted for both taxa in Provence, with in particular a clear decrease in asymmetry, an increase in the intraspecific variability, and an interspecific convergence of karyotypes. These changes suggest a drift that has facilitated various crosses, and has been amplified through hybridization/introgression. Furthermore, several morphological and karyological transgressive characters have been found in the two invasive taxa. These results stress the important role and the rapidity of karyological changes in invasive processes.