Do the introductions by botanical gardens facilitate the invasion of Solidago canadensis (Asterceae) in China?

Many invasive plants have long been suspected of firstly being introduced and cultivated by a local botanical garden or nursery and then escaping into the field after adapting to the novel environment. The role of botanical gardens in the spread of invasive plants has not yet been explored experimentally. In this article, we studied the possible roles of two botanical gardens in the spread of invasive Solidago canadensis (Asterceae) in China by analysing genetic relationships of invasive and native (United States) populations with intersimple sequence repeats markers. Our results showed a high genetic variation (mean He = 0.292) and a large proportion of genetic variation (85.6%) residing within populations. Solidago canadensis was possibly introduced firstly into eastern China. The plants from Lushan Botanical Garden showed distant genetic distance from all of the other populations, suggesting that this botanical garden had little effect on the invasion of S. canadensis. Populations from Wuhan Botanical Garden in central China, however, showed close genetic relationships with local populations and populations in west and south‐west China, suggesting gene exchange between these populations. Thus, risk assessment is critical for plant introduction and conservation, as introductions of alien plants by botanical gardens may facilitate plant invasions, while plants conserved in botanical gardens may be at risk by surrounding plant invasions.     

Genetic variation of the invasive Campuloclinium macrocephalum,Asteraceae in South Africa,inferred from molecular markers

Campuloclinium macrocephalum is native to Central and South America, but is a highly invasive weed in South Africa, where it is commonly known as the ‘pompom weed’. It is targeted for biological control, the success of which will depend on host specificity and biotype compatibility to its full genetic diversity in South Africa. We investigated the genetic diversity and phylogeography of 52 specimens from across South Africa, 14 from Argentina and three from Brazil using nuclear ribosomal ITS regions. We further explored the AFLP marker diversity in 54 South African, 25 Argentine and three Brazilian specimens. Maximum parsimony analysis of the ITS sequence data produced an unresolved phylogeny. However, three haplotypes were recognised via network analysis. All South African, one Brazilian and all bar one of the Argentine individuals shared a single haplotype. AFLP analyses generated two genetic clusters with a low net nucleotide distance of 0.115 and further revealed that most plants were a mixture of alleles from these two genetic clusters. Although there was a significant genetic variation among the populations, genetic differentiation and mean heterozygosity were low, suggesting that clonal reproduction may be occurring. The current South African populations may therefore be clonal products with different proportions of genetic admixture introduced more than once. The original point of entry appears to be Gauteng, South Africa. Long‐distance dispersal appears to have played a major role in its spread across South Africa. Candidate Argentine biological control agents should therefore be effective on C. macrocephalum in South Africa.     

Genetic structuring and diversity patterns along rivers – local invasion history of Ambrosia artemisiifolia (Asteraceae) along the Danube River in Vienna (Austria) shows non‐linear pattern

Ambrosia artemisiifolia is an annual weed from North America that nowadays is invasive in many countries worldwide. In Austria, numerous populations of A. artemisiifolia are located along the Danube River, especially along the ‘New Danube’ (Vienna). This area is characterised by ruderal and riparian sites, which are regularly flooded. To better understand the spread of A. artemisiifolia and its colonising behaviour along the Danube River, we analysed genetic structure and diversity based on 23 populations linearly arranged along the Viennese Danube riverbed and upstream, utilising the Amplified Fragment Length Polymorphism (AFLP) fingerprint method. We generated 284 polymorphic AFLP markers across 446 A. artemisiifolia plants. The genetic diversity within populations was higher (HW = 0.091) than among populations (HB = 0.007). This result indicates A. artemisiifolia introductions from similar mixtures of sources or spread from a single already mixed introduction. Within our local setting, we were unable to identify neither source or sink populations nor an obvious linear genetic structuring. Genetic among‐population differentiation was low to moderate (amova ‐derived F_ST = 0.124). Lack of geographical structuring is indicative of highly dynamic gene flow, which is further supported by the absence of an isolation‐by‐distance pattern. Multiple introductions and non‐directional gene flow are most likely promoted by anthropogenic disturbance and human‐mediated dispersal. Our results demonstrate the ability and speed of A. artemisiifolia to settle in newly disturbed areas and the difficulties to predict invasion directions, as downstream river dispersal was negligible.     

Invasion process and potential spread of Amaranthus retroflexus in China

Amaranthus retroflexus, a troublesome agricultural weed native to North America, has expanded its distribution in large areas of China since its introduction around 1905. Geographical interpretation of changes in their distribution ranges could provide valuable insights on its spatiotemporal invasion patterns and could be used to predict the extent of its future spread. Based on compiled historical distribution occurrences of A. retroflexus in North American and Chinese ranges, invaded ecological niche models for three hypothetical invasion stages were developed. Native models on the basis of all available records within the North American range were also generated for reciprocal comparison with the invaded model. Climate similarity between native and invasive ranges was also investigated. Amaranthus retroflexus has exhibited a rapid and large range expansion after about a 50‐year lag, especially in central and western China. It established a relative stable distribution in the 1960s and has been undergoing a more continuous westward expansion since then. Presently A. retroflexus has not yet reached full occupancy of suitable habitats in China. The results highlight prioritising habitats in south‐western China for monitoring and control to prevent its further spread.     

Genetic diversity of Pueraria lobata (kudzu) and closely related taxa as revealed by inter-simple sequence repeat analysis

Kudzu (Pueraria lobata) is a noxious weed infesting some areas of the USA. Knowledge of its genetic variation in both native and invasive areas can lead to effective biological control measures. Inter-simple sequence repeat (ISSR) variations were studied in P. lobata and its four closely related congeneric species (P. edulis, P. montana, P. phaseoloides and P. thomsoni). ISSR results allowed a clear separation of these five species. For P. lobata, 108 plants from China and USA were analysed. The samples from the US were genetically closer to Chinese P. lobata populations than to other congeneric populations. High genetic differentiation was found for P. lobata, P. montana and P. thomsoni in Chinese samples. High genetic diversity and low population differentiation was found in P. lobata samples of the US. This supports the hypothesis of multiple introductions into the USA from different sources in Japan or China, followed by subsequent gene exchange and recombination.     

Assessing wind and mammals as seed dispersal vectors in an invasive legume

While some plants have modified seed structures to facilitate dispersal, many lack such specialised adaptations, making their mode of dispersal unclear. This can be particularly problematic for predicting shifts in species ranges or tracking the spread of invasive plants. As an example, the seed size and shape of the invasive legume, Lespedeza cuneata, suggest that wind and attachment to animals are not important for dispersal, yet populations can spread surprising distances within a few years. Using a series of experiments conducted in the laboratory and field, we tested wind and mammal fur as mediators of seed dispersal. To test wind dispersal, seed traps were arranged radially around a patch of L. cuneata and seeds were collected following dispersal. Attachment to mammal fur was tested by fitting pelts of a deer, coyote and raccoon to artificial torsos and determining seed retention in both the field and the laboratory. Laboratory trials also examined the influence of wet versus dry conditions. Our results showed that wind direction strongly influenced dispersal distance and seeds were readily dispersed by mammal fur. The number of seeds retained was species specific, depending on fur depth and mammal size, with seed retention increasing under wet conditions. Together, these results suggest that both wind and mammal fur contribute to the movement of L. cuneata across grasslands. Consequently, both dispersal vectors should be considered when designing and implementing control strategies.     

Are road verges corridors for weed invasion? Insights from the fine‐scale spatial genetic structure of Raphanus raphanistrum

Raphanus raphanistrum (Brassicaceae) is considered amongst the world's worst agricultural weeds. We address critical issues in its management by studying the pathway of colonisation at local scales. For this, we assessed the small‐scale spatial genetic structure of 231 samples collected from three different sites across the Cape Floristic Region, South Africa, using 11 nuclear microsatellite markers. Although natural pollen and seed dispersal were expected to be restricted, we found no significant relationship between genetic and geographical distance within sites. Instead, our results suggest that R. raphanistrum had colonised new habitats via jump dispersal, rather than through natural diffusive dispersal at local scales. We did not find evidence for road verges as dispersal corridors, as evidenced by a lack of isolation‐by‐distance at local scales. Instead, the absence of spatial genetic structure suggests that R. raphanistrum had rapidly spread throughout its current range, possibly facilitated by human‐mediated actions. Management plans addressing containment or suppression of the weedy species R. raphanistrum (and possibly other weedy species) should take the high degree of connectivity between distant geographical localities into account.     

Genetic analysis of Phytophthora sojae populations in Fujian, China

Phytophthora sojae is a destructive soilborne pathogen causing seedling damping‐off and root rot of soybean (Glycine max). The goal of this study was to determine the genetic structure of P. sojae populations in Fujian, China. Nine microsatellite markers were used to investigate the genetic variation in 19 P. sojae populations, sampled from Fujian Province and northeastern China (Jilin and Heilongjiang Provinces) between 2002 and 2013. Overall, a low genetic diversity, Hardy–Weinberg disequilibrium, and an index (an index of association) that was significantly different from zero were detected in populations; these results were consistent with self‐fertilization and clonal modes of reproduction for this pathogen. However, using Bayesian Markov chain Monte Carlo approach, principal component analysis and neighbour joining (NJ) algorithm, the Fujian P. sojae populations clustered into three distinct groups, one of which included most isolates of the northeast populations. What is more, significant estimates of pairwise fixation indices (F_ST) were detected between most populations, especially in different clusters. It is hypothesized that the cropping system used, the limited dispersal ability, and human‐mediated gene flow may account for the observed genetic structure of P. sojae populations in Fujian, China. In addition, a high virulence frequency of the pathogen on different cultivars carrying known major R genes for resistance, and a rapid increase in virulence frequency, indicated that these major R genes should not be used to manage seedling damping‐off and root rot diseases of soybean (Glycine max).     

Genetic analysis of the population structure of the rice false smut fungus,Villosiclava virens,in China using microsatellite markers mined from a genome assembly

Studies on population genetics of Villosiclava virens are limited because of the lack of polymorphic markers. Based on a draft genome sequence of isolate HWD‐2 produced in this study, 20 of 403 potential simple sequence repeats (SSR) loci showed polymorphisms in preliminary screening using eight diverse V. virens isolates. Among polymorphic loci, most of them with tetra‐ to hexanucleotide unit motifs showed higher levels of polymorphism than loci with smaller motifs. After testing with 20 polymorphic SSR markers, the 87 isolates of V. virens from eight populations in China showed a high level of genetic diversity, with each as a unique haplotype. This differs from some previous findings showing little to no genetic variation based on random amplified polymorphic DNA and amplified fragment length polymorphism analyses. Among eight populations from major rice production areas of China, the population from Guangxi province in south China showed the highest levels of polymorphism, which led to the speculation that it might be closer to the centre of origin of this pathogen. The northern, central and eastern populations (Jilin, Liaoning, Hubei, Hunan, Jiangxi and Zhejiang), when considered together as a group, showed significant molecular variation compared to the southern populations (Fujian and Guangxi) (Φ_PT = 0·043, P = 0·037). A significant relationship (Mantel test, P = 0·027) but with low correlation (R² = 0·23) was also found between geographic distance and genetic distance. The 20 polymorphic SSR primer pairs designed in this study provide a tool for further research on the population diversity of this emerging fungal pathogen of rice.     

Genetic diversity and structure of Hemileia vastatrix populations on Coffea spp.

Coffee leaf rust is the most limiting disease for coffee cultivation in Brazil. Despite its importance, relatively little is known about the genetic diversity of Hemileia vastatrix, the rust causal agent. In this work, the DNA from 112 monopustule isolates from different geographic locations and coffee genotypes were analysed by amplified fragment length polymorphisms (AFLP). The objectives were to assess the influence of the host and geographic origin on the diversity and population differentiation in H. vastatrix. The fungal population showed a low level of genotypic diversity. Gene diversity (h) was 0·027 and the hypothesis of random mating in the total population was rejected, but evidence for recombination was found for two subpopulations (São Paulo and Paraná). The analysis of molecular variance revealed that 90% of the genetic distribution of the pathogen occurs among isolates within the subpopulation (states or host of origin). There was no correlation between geographic and genetic distance (r = ?0·024, P = 0·74), which together with the high number of migrants and the low degree of differentiation in populations of H. vastatrix, is consistent with the fact that the inoculum is probably easily dispersed by wind over long distances, allowing dispersal of the pathogen among coffee growing areas in Brazil. Therefore, it is difficult to predict the durability of resistant sources to coffee rust. The recommendation for the breeding programmes is thus to incorporate multigenic resistance as a control strategy.     

Genetic diversity within and between sulfonylurea‐resistant and susceptible populations of Schoenoplectus juncoides in Japan

To reveal the effects of herbicide selection on genetic diversity in the outcrossing weed species Schoenoplectus juncoides, six sulfonylurea‐resistant (SU‐R) and eight sulfonylurea‐susceptible (SU‐S) populations were analysed using 40 polymorphic inter‐simple sequence repeat loci. The plants were collected from three widely separated regions: the Tohoku, Kanto and Kyushu districts of Japan. Genetic diversity values (Nei's gene diversity, h) within each SU‐S population ranged from h = 0.125 to h = 0.235. The average genetic diversity within the SU‐S populations was H_S = 0.161, and the total genetic diversity was H_T = 0.271. Although the H_S of the SU‐R populations (0.051) was lower than that of the SU‐S populations, the H_T of the SU‐R populations (0.202) was comparable with that of the SU‐S populations. Most of the genetic variation was found within the region for both the SU‐S and SU‐R populations (88% of the genetic variation respectively). Two of the SU‐R populations showed relatively high genetic diversity (h = 0.117 and 0.161), which were comparable with those of the SU‐S populations. In contrast, the genetic diversity within four SU‐R populations was much lower (from h = 0 to 0.018) than in the SU‐S populations. The results suggest that selection by sulfonylurea herbicides has decreased genetic diversity within some SU‐R populations of S. juncoides. The different level of genetic diversity in the SU‐R populations is most likely due to different levels of inbreeding in the populations.     

Vegetative propagation of Solidago canadensis – do fragment size and burial depth matter?

Clonal species may benefit from human disturbance because their vegetative fragments may be distributed via soil. Solidago canadensis is an invasive rhizomatous perennial frequently found in ruderal environments. When creating new infrastructure, digging and cutting are two main factors that may influence the spread of S. canadensis into new areas. To have a better understanding of the invasive potential of S. canadensis, we investigated whether S. canadensis was able to survive and grow from stem cuttings as well as from rhizomes. Rhizomes and cuttings were collected from three populations in Eastern Norway. The rhizomes and cuttings were planted in a pot experiment to assess their vegetative ability to propagate. Rhizome fragments (5 and 10 cm long) were buried at 0.5, 10 and 30 cm depths. The cuttings were planted as 15 cm stems, with the bottom 5 cm pushed into the soil. The results showed that rhizome length did not have an effect on survival. Although some sprouting occurred at all burial depths, increasing depth had a negative effect on rhizome survival. In general, development of the cuttings was good, but there were differences between population performance and survival. These results imply that care must be taken when (i) constructing new sites, because digging and transport of soil masses may spread S. canadensis into new areas by rhizomes or cuttings, and (ii) mowing road verges and other ruderal areas to prevent the spread of stem cuttings from one area to another.     

Pathogenic and molecular diversity in highly virulent populations of the parasitic weed Orobanche cumana (sunflower broomrape) from Europe

The parasitic weed Orobanche cumana (sunflower broomrape) constrains sunflower production in eastern and southern Europe and in the Middle East. Although genetic resistance is the most effective control method, new parasite races evolve overcoming sunflower resistance. In this work, highly virulent populations of O. cumana were analysed for pathogenicity and genetic diversity. The virulence of 11 populations from Hungary, Romania, Spain and Turkey was assessed and compared after infection of sunflower inbred lines to differentiate races of the parasite under glasshouse conditions. Molecular diversity among and within 27 parasite populations was studied by RAPD‐PCR, UPGMA and amova analyses. Highly virulent race F was identified in Hungary, Spain and Turkey. The most virulent race (G) was also found in Turkey. The molecular analysis among highly virulent populations of O. cumana identified four molecular clusters, respectively, grouping populations from Central Spain, Hungary, South Spain and Turkey. The genetic homogeneity within parasite populations was confirmed, since no molecular divergences were found within them. This work constitutes the first geographical study of O. cumana together with pathogenicity and molecular traits inherent to each geographical group, and provides useful information for possible phylogenetic analyses of O. cumana. In addition, molecular markers associated with geographical origin could be developed and used as diagnostic tools to track new broomrape introductions into areas free of virulent races where they might represent a threat to sunflower production.     

Genetic structure of Verticillium dahliae isolates infecting olive trees in Tunisia using AFLP,pathogenicity and PCR markers

Since 2006, verticillium wilt of olive induced by Verticillium dahliae has caused considerable economic losses in olive orchards in Tunisia. The genetic structure of V. dahliae isolates collected from different olive growing regions was investigated using virulence tests, vegetative compatibility grouping (VCG) and amplified fragment length polymorphism (AFLP) analyses. In total, 42 isolates of V. dahliae from diseased olive trees were tested. Cluster analysis and principal coordinate analysis revealed that geographic origin was the main factor determining the genetic structure of V. dahliae populations and both methods indicated a genetic separation between the central and coastal isolates. Isolates were divided into two major groups: the AFLP‐I group included all isolates from Sidi Bouzid, Kairouan, Kasserine and Sfax (centre of the country) and the AFLP‐II group included isolates from Monastir, Zaghouane, Sousse, Mahdia (coastal region), and two isolates from Sfax. Analysis of the molecular variance (amova ) indicated a significant level of genetic differentiation among (76%) and within (23%) the two populations. Analyses of both the defoliating (D) and non‐defoliating (ND) pathotypes and VCG markers indicated that most of the isolates belong to VCG 2A and 4B/ND pathotype. The disease severity was highly variable among the isolates tested (P < 0·05) with no evidence of association between aggressiveness and geographical origin of the isolates. Overall, results of this study revealed a clear association between the genetic diversity of the isolates and their geographic origin, but not between genetic diversity and virulence patterns.     

Genetic diversity of giant reed ( A rundo donax) in Australia

The perennial grass, Arundo donax, has shown potential as a promising biomass crop. However, it has become invasive in a number of areas and declared a noxious weed in some jurisdictions, making proposals to grow A. donax for commercial use in Australia controversial. Evidence of asexual reproduction and the presence of a single genetic clone in Australia was investigated, as such characteristics would indicate a limited risk of escape and invasion. Using amplified fragment length polymorphism markers, the genetic diversity of 218 A. donax samples from across Australia was examined. The samples were found to separate into two distinct genetic groups, or clades. There was only a small amount of genetic diversity within a clade (0.9 and 1.5%). However, there was a larger difference between the clades of 19.8%, suggesting the presence of two distinct A. donax genotypes in Australia. The low level of genetic variation in Australian A. donax that was found in this study indicates that spread is essentially by vegetative means and suggests that if grown in areas where it is separated from natural water dispersal events, A. donax poses a low risk of becoming invasive.     

High genotype diversity and lack of isolation by distance in the Alternaria solani populations from China

A genomic library was used to develop seven SSR markers for studying the population genetics of Alternaria solani, a pathogenic fungus causing early blight disease of potato and tomato worldwide. Population genetic analysis of 268 isolates of A. solani sampled from four locations, each representing one of four potato production systems in China, indicates that these SSR markers are moderately diverse, selectively neutral and possibly unlinked. Population genetic analysis also indicated that genetic variation of A. solani in China is high. About 2/3 of 123 genotypes were detected only once and genotype diversity measured by the standardized Shannon index ranged between 0·82 and 0·92 in the populations. Although clones were detected in multiple populations separated by thousands of kilometres, random association among SSR loci was found in half of the populations assayed. On average, nearly six copies of genetic material were exchanged among these populations each generation and no isolation by distance was detected. It is hypothesized that the joint effects of cryptic sexual reproduction and human‐mediated gene flow may account for the observed population genetic structure of A. solani in China.     

Evolution and spread of glyphosate resistance in Conyza bonariensis in California and a comparison with closely related Conyza canadensis

Glyphosate‐resistant weeds are an increasing problem in perennial cropping systems in the Central Valley of California, USA. To elucidate the evolutionary origins and spatial spread of resistance, we investigated the geographical distribution of glyphosate resistance and the population genetic diversity and structure of Conyza bonariensis and compared the results with previously studied C. canadensis. Thirty‐five populations from orchards and vineyards across the Central Valley were sampled. Population genetic structure was assessed using microsatellite markers. Population‐level resistance was assessed in glasshouse screening of plants grown from field‐collected seed. Bayesian clustering and analyses of multilocus genotypes indicated multiple origins of resistance, as observed in C. canadensis. Pairwise F_ST analysis detected spatial spread of resistance in the south of the Central Valley, also similar to C. canadensis. The results strongly indicate that the southern valley was an environment markedly more suitable than the northern valley for resistance spread and that spread in Conyza species was driven by increased uniformity of strong selection in the southern valley, due to recent regulation on herbicides other than glyphosate. Accordingly, resistant C. canadensis individuals occurred at high frequencies only in the southern valley, but interestingly resistant C. bonariensis occurred at high frequencies throughout the valley. Expression of resistance showed varying degrees of plasticity in C. bonariensis. The lower selfing rate and substantially greater genotypic diversity in C. bonariensis, relative to C. canadensis, indicate greater evolutionary potential over shorter time periods. Interspecific hybridisation was detected, but its role in resistance evolution remains unclear.     

Populations of Ceratocystis fimbriata on Colocasia esculenta and other hosts in the Mata Atlântica region in Brazil

Ceratocystis fimbriata is native to Brazil, where it is able to cause serious diseases on numerous hosts, especially on non‐native plants. Because C. fimbriata is soilborne and not wind dispersed, highly differentiated populations are found in different regions of Brazil. The present study compared populations of C. fimbriata on taro, mango, eucalyptus and kiwifruit from the coastal Mata Atlântica region with native populations of the fungus from the Cerrado‐transition region in Brazil by using 14 SSR markers and DNA sequences of ITS and mating type genes. Microsatellite and phylogenetic analyses were performed to test the hypothesis that populations on different hosts from the Mata Atlântica region are related to each other and are native to the region. The ITS sequences varied greatly among the taro isolates, with six sequences identified, from which two had not been previously reported. For mating type genes, four sequences were identified among the isolates on taro, mango, eucalyptus and kiwifruit. Phylogenetic analyses showed that Mata Atlântica populations formed a monophyletic group distinct from Cerrado‐transition region populations, although earlier studies had shown that isolates from the two regions are interfertile and are considered as a single biological species. Microsatellite analysis revealed low gene diversity for each of the three Mata Atlântica populations on taro, mango and kiwifruit, suggesting that these populations had gone through genetic bottlenecks, probably by dispersal of select genotypes in vegetative propagation material. Also, microsatellite markers showed that two microsatellite genotypes from taro are widely spread in Brazil, probably by infected corms.     

Miscanthus x giganteus crop fields hide a genotype of the invasive M. sacchariflorus

Despite their putative invasive potential, Miscanthus taxa have been increasingly viewed as promising crops for bioenergy production. Miscanthus x giganteus is cultivated worldwide due to its high productivity, and its non‐spreading growth limits its risk of invasion. However, the genetic uniformity of its unique cultivated clone has recently been questioned. We used morphometry, nuclear microsatellites, AFLP fingerprints and nrDNA ITS barcoding to assess Miscanthus genotypes cultivated in eastern France. As expected, the globally cultivated clone was the main morphotype and genotype of our samplings; however, we also identified a second genotype, assigned to M. sacchariflorus, in the sampled fields. Morphological differentiation partially overlaps genotypic assignation, probably due to disturbance at the crop edges. We also show that Miscanthus taxa are misidentified in commercial nurseries. Our study confirms previous reports of the occurrence of other accessions in fields of M. x giganteus. These taxonomic confusions limit the certainty that a single clone of M. x giganteus is cultivated. In this context, M. x giganteus crop fields may actually promote the dispersal of invasive Miscanthus taxa.     

Genetic structure of the invasive Chromolaena odorata in China

Chromolaena odorata is one of the world's worst tropical weeds, and it can be found in Guangdong, Guangxi, Hainan and Yunnan provinces in China. Genetic variation of 27 C. odorata populations sampled from across its invasive range in China was investigated using inter‐simple sequence repeats (ISSR) analysis. All populations exhibited low levels of genetic variation. On average, 2.35% of the ISSR loci were polymorphic, total genetic diversity (H_T) was 0.0406 and Shannon's information index (H_sp) was 0.0623. High genetic identities (I) between populations (0.9687 ± 0.01204) indicated low genetic differentiation. A frequent founder effect was interpreted as the main cause of the genetic structure observed in C. odorata.