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Christoph Chucholl Predicting the risk of introduction and establishment of an exotic aquarium animal in Europe: insights from one decade of Marmorkrebs Crustacea, Astacida, Cambaridae releases pp Download article Open Access. The presence of the North American Marmorkrebs Procambarus fallax f. Since at least 25 independent introduction events have been confirmed, of which some have eventually resulted in established populations. This study links a maximum-entropy model that forecasts the probability of Marmorkrebs introduction based on socio-economic predictors to an updated species distribution model based on environmental predictors in order to explore the risk of further Marmorkrebs establishment in Europe. In line with related research, the probability of Marmorkrebs release was largely affected by gross domestic product and human population density, i. While considerable parts of Europe were predicted as potentially suitable for establishment, high probabilities of introduction were forecasted in much smaller geographic areas. The consensus map of the model predictions suggests that introduction and subsequent establishment of Marmorkrebs is likely to occur in much of Germany, the Benelux countries, England, Italy, and areas of high human population density throughout France and Spain, as well as parts of southernmost Scandinavia and Southeastern Europe. Monitoring trades of Marmorkrebs in these high-risk regions is recommended and implications for proactive measures are discussed, including the need for consistent trade regulations at the EU level. Lucas R. Nathan, Christopher L. Mahon An assessment of angler education and bait trade regulations to prevent invasive species introductions in the Laurentian Great Lakes pp The commercial bait trade is one pathway for aquatic invasive species AIS introductions, as non-target bait species can be accidentally sold to anglers who either inadvertently or intentionally release them while fishing or as excess bait. Prevention of AIS introductions via the bait trade requires a two-tiered management approach, which includes both regulation of the bait industry and angler education. Retail bait shops may offer opportunities for public education regarding AIS, however it is unknown how often shops are targeted for such purposes and how viable this method of angler education is on a temporal scale. The goals of this research were to 1 quantify the current distribution of AIS signage in retail bait shops in the Great Lakes region and 2 estimate the long term viability of using retail bait shops as platform for angler education. Additionally, we present an up-to-date summary of bait industry regulations across the Great Lakes jurisdictions. The presented summary of bait regulations for Great Lakes jurisdictions indicates multiple discrepancies across the region, which may hinder successful management strategies. Future management goals should consider additional methods of angler education and coordinating regulations across the Great Lakes to improve upon AIS prevention. Daniel Clements, Tony M. Dugdale, Kym L. Butler and Trevor D. Hunt Management of aquatic alligator weed Alternanthera philoxeroides in an early stage of invasion pp Alligator weed Alternanthera philoxeroides Mart. It has invaded at least 14 countries and is difficult to control. The present study investigates the effectiveness of herbicides and physical removal in eliminating patches of aquatic alligator weed in an early stage of invasion. The presence of surfactant did not improve metsulfuron-methyl efficacy. Field studies were then carried out to evaluate the effectiveness of repeated physical removal and repeated applications of chosen herbicides to eliminate patches of aquatic alligator weed in an early stage of invasion of two urban streams in Melbourne, Australia. Glyphosate and metsulfuron-methyl without a surfactant were applied to patches of aquatic alligator weed in a best practice regime, consisting of up to three applications per year for up to five consecutive years. No alligator weed remained after two years of the herbicide application regime for patches treated with metsulfuron-methyl, while for glyphosate alligator weed remained in only one of 18 patches after three years. This study demonstrates that the management methods utilised are capable of eliminating patches of aquatic alligator weed in an early stage of invasion in two to three years. The red swamp crayfish Procambarus clarkii has been widely introduced into aquatic systems all around the world, triggering multiple impacts. Water persistence, salinity, and substrate can condition the presence of the species in the wild. Once declared a protected area, managers assumed control of the maintenance of the park and opened new water points as a wildlife management tool. Crayfish occurrence was significantly related to the presence of seasonal streams connecting the ponds with refuge areas and to the presence of artificial water points within the pond. The probability of occurrence of red swamp crayfish in a pond without seasonal streams was more than 5 times lower than that in a pond with seasonal streams. The probability of crayfish occurrence in ponds without artificial water points was 2. We review the advantages and disadvantages of water points as a management tool. Gary C. Matlock Temporal trends in non-native fishes established in the continental United States pp Management of non-native fish species is informed by monitoring their temporal and spatial distribution. There are few published analyses of temporal patterns of established non-native fish species. As of the AFS list, there were 66 non-native species listed as established in the continental U. All of the species, except two, established before persisted in The number of species added to each subsequent list increased throughout the period. Eight species were added from to , 14 from to , 18 from to , and 5 from to The number of listed non-native families increased linearly from 8 to 19 during the same period. Converting the trend in the number of non-native species into the number of invasive species is difficult because of lack of available impact information and the dependency on human perception and value. The silver-cheeked toadfish Lagocephalus sceleratus is one of the most recent invaders in the Mediterranean Sea and a serious risk for public health. In June , an awareness campaign to disseminate information related with this toxic species was launched by the Tunisian Ministry of Agriculture. Whilst providing a step toward the development of an appropriate early warning system, this case highlights a remarkable geographical extension of L. We illustrate how the first step in the process of effectively managing the risks posed by an invasive species is to engage and communicate with the public. James M. Long, Yu Liang, Daniel E. Shoup, Andrew R. Dzialowski and Joseph R. Bidwell GIS-based rapid-assessment of bighead carp Hypophthalmichthys nobilis Richardson, suitability in reservoirs pp Broad-scale niche models are good for examining the potential for invasive species occurrences, but can fall short in providing managers with site-specific locations for monitoring. Using Oklahoma as an example, where invasive bighead carp Hypophthalmichthys nobilis are established in certain reservoirs, but predicted to be widely distributed based on broad-scale niche models, we cast bighead carp reproductive ecology in a site-specific geospatial framework to determine their potential establishment in additional reservoirs. Moreover, this method was rapid and identified sites that could be prioritized for increased study or scrutiny. Our results highlight the importance of considering the environmental characteristics of individual sites, which is often the level at which management efforts are implemented when assessing susceptibility to invasion. Danielle M. Haak, Bruce J. Stephen, Robert A. Kill, Nicholas A. Smeenk, Craig R. Allen and Kevin L. Pope Toxicity of copper sulfate and rotenone to Chinese mystery snail Bellamya chinensis pp The Chinese mystery snail Bellamya chinensis is a freshwater snail native to Southeast Asia, Japan, and Russia and is currently classified as an invasive species in at least 27 states in the USA. The species tolerates a wide range of environmental conditions, making management of established populations difficult. We tested the efficacy of two traditional chemical treatments, rotenone and copper sulfate, on the elimination of adult Chinese mystery snails in laboratory experiments. Neither rotenone nor copper sulfate effectively killed adult Chinese mystery snails in laboratory experiments, most likely due to their relatively large size, thick shell, and operculum. Therefore, it appears that populations will be very difficult to control once established, and management should focus on preventing additional spread or introductions of this species. Osariyekemwen O. Uyi, Frank Ekhator, Celestine E. Ikuenobe, Temitope I. Borokini, Emmanuel I. Aigbokhan, Ikponmwosa N. Egbon, Abiodun R. Adebayo, Igho B. Igbinosa, Celestina O. Okeke, Etinosa O. Igbinosa and Gift A. Omokhua Chromolaena odorata invasion in Nigeria: A case for coordinated biological control pp Chromolaena odorata L. King and Robinson Asteraceae: Eupatorieae is an invasive perennial weedy scrambling shrub of neotropical origin, widely acknowledged as a major economic and ecological burden to many tropical and subtropical regions of the world including Nigeria. Here, we examine the invasion and management of C. We searched major institutional libraries in Nigeria and carried out extensive research of historical records using different data base platforms, including Google Scholar, Science Direct, ISI web of Science, SciFinders and Scopus. Apart from the biological invasive characteristics of C. The current distribution and infestation levels of this invasive weed in Nigeria are mapped. The current methods of control and the failed attempt made by the Nigerian government to eradicate the weed between the late s and s are discussed. We argue that even in the face of the usefulness of C. Matthew R. First, Stephanie H. Robbins-Wamsley, Scott C. Riley, Jacqueline I. Fisher, Joseph P. Smith and Lisa A. Rigorous evaluation of ballast water management systems BMWSs at land-based test facilities requires that water used in testing meets minimum concentrations of dissolved and particulate material, for example, using the criteria in the U. In this study, we evaluated materials used to supplement dissolved organic matter DOM , particulate organic matter POM , and mineral matter MM used to achieve challenge water criteria. Additionally, the response of ambient organisms to these additives was measured in mesocosm experiments, in which changes in organism concentrations were measured after a 5-d holding time. A field trial was carried out to test the respective efficacy of three different mechanical dredge methods at reducing Corbicula clam numbers by estimating changes in abundance and biomass immediately following dredging. Three sites which supported different population density and biomass levels within the tidal section of the River Barrow were subject to each of the dredge methods. The methodology and results indicate that, while dredging can achieve a large reduction in Corbicula population numbers, further research is required before this can be considered as a management tool for control of Asian clams. Published in November Contents Research articles Christoph Chucholl Predicting the risk of introduction and establishment of an exotic aquarium animal in Europe: insights from one decade of Marmorkrebs Crustacea, Astacida, Cambaridae releases pp Abstract Download article Open Access The presence of the North American Marmorkrebs Procambarus fallax f. Mahon An assessment of angler education and bait trade regulations to prevent invasive species introductions in the Laurentian Great Lakes pp Abstract Download article Open Access The commercial bait trade is one pathway for aquatic invasive species AIS introductions, as non-target bait species can be accidentally sold to anglers who either inadvertently or intentionally release them while fishing or as excess bait. Hunt Management of aquatic alligator weed Alternanthera philoxeroides in an early stage of invasion pp Abstract Download article Open Access Alligator weed Alternanthera philoxeroides Mart. Matlock Temporal trends in non-native fishes established in the continental United States pp Abstract Download article Open Access Management of non-native fish species is informed by monitoring their temporal and spatial distribution. Bidwell GIS-based rapid-assessment of bighead carp Hypophthalmichthys nobilis Richardson, suitability in reservoirs pp Abstract Download article Open Access Broad-scale niche models are good for examining the potential for invasive species occurrences, but can fall short in providing managers with site-specific locations for monitoring. Pope Toxicity of copper sulfate and rotenone to Chinese mystery snail Bellamya chinensis pp Abstract Download article Open Access The Chinese mystery snail Bellamya chinensis is a freshwater snail native to Southeast Asia, Japan, and Russia and is currently classified as an invasive species in at least 27 states in the USA. LOG IN. Events Calendar. Decision Support System. Black Sea Region. Baltic Sea Region. NE Baltic Sea. Editorial Board. Current Issue. Manuscript Submission. Volume 14, Issue 1 March Volume 14, Issue 2 June Volume 14, Issue 3 September Volume 14, Issue 4 November Volume 15, Issue 1 March Volume 15, Issue 2 June Volume 15, Issue 3 September Published in November Contents. Rory Sheehan, Joseph M.

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Carla Lambertini, Brian K. Two Phragmites australis taxa are recognized in Europe: P. Another taxonomic group in the Mediterranean is Phragmites frutescens. European genotypes are diverse genetically, cytologically and morphologically, and are related to African, Asiatic and American genotypes. We analysed chloroplast variation with parsimony and genetic distance methods, and compared it with that of nuclear amplified fragment length polymorphism and microsatellites. We also investigated the phenological pattern of genotypes collected worldwide in a common garden in Denmark. The European P. However, African and North American genotypes are differentiating from the European genotypes. Mediterranean P. Phragmites frutescens shares the cpDNA sequences with both these tropical species. Phenological data confirmed possible gene flow within the temperate region of Europe, whereas the Mediterranean genotypes did not set inflorescences in Denmark, suggesting reproductive barriers between temperate and Mediterranean P. European P. Further research is needed to understand the implications of long-distance dispersal at the population level. The genus Phragmites Adans. Poaceae is cosmopolitan with a distribution ranging from the tropics to cold temperate areas of both hemispheres Clevering and Lissner The Phragmites species are tall up to 6 m , perennial grasses predominantly found in littoral zones of lakes, along rivers and in shallow freshwater and brackish wetlands Brix At present, five species are recognized, of which Phragmites australis Cav. Trin ex Steudel is the most widely distributed Clevering and Lissner There is, however, morphological and cytological variation within Phragmites species, and in particular within the cosmopolitan P. In Europe, two P. The name P. The Flora Europeae Tutin et al. The recent recognition of Phragmites frutescens H. The morphological similarities of P. Lambertini et al. The subspecific rank for P. Phragmites australis is the former Phragmites communis Trin. Conert merged 64 taxa into P. All varieties were present in Europe, and most of them in a limited geographic area, such as Germany. Even though Conert's classification system has been abandoned, it reflects very well the often gradual morphological variation in size, colour and leaf structure present in Europe, sometimes observable even among adjacent stands Rolletschek et al. In addition to P. Several genetic studies at the population level conducted throughout Europe have consistently shown that genetic variance is distributed within, rather than among, populations, and that populations can be genetically very different from each other even at a local scale Lambertini et al. However, isolation-by-distance patterns have failed to be detected Lambertini et al. Phylogeographically, European Phragmites belongs to a group that extends throughout Eurasia and North America, as well as in the Southern Hemisphere in Africa and Australia Saltonstall ; Lambertini et al. The lack of structure within the European group, as opposed to other geographically distinct P. When relationships among genotypes are analysed at the continental scale, the idea of a single population of randomly mating individuals seems to work well for European Phragmites Lambertini et al. However, genetic distances among adjacent stands may be high and comparable to those observed among stands hundreds, if not thousands, of kilometres apart Lambertini et al. The cryptic invasion of a Euroasiatic cp-lineage in North America, described as haplotype M Saltonstall , shows that a single maternal lineage can establish in a variety of habitats and populate an entire continent. In contrast to many invasive clonal species Geng et al. Genetic diversity within the Euroasiatic lineage appears therefore to be a key factor explaining its broad ecological amplitude. In the European range, P. In the Gulf Coast of the USA, however, the nuclear alleles of the Euroasiatic lineage are introgressed in another distinctive cp-lineage, known as haplotype I Saltonstall , which has recently been revealed to be an interspecific hybrid between P. The first aim of this study was to analyse chloroplast DNA cpDNA diversity in Europe and the phylogeographic relationships of European haplotypes at the global scale. The second aim was to identify the current borders of the European gene pool in order to determine the range within which gene flow is likely to be occurring, and thus understand the intercontinental relationships of European Phragmites. Given the broad distribution of European-related P. Differently from previous studies we compared the global pattern of phylogeographic relationships inferred by cpDNA with that of nuclear DNA. We also examined phenology in a common garden in Denmark of Phragmites genotypes from all over the world, in order to have a model system in which to observe the implications of long-distance interrelationships and their congruence with molecular markers. The hypothesis behind the common garden was that genotypes releasing pollen and seeds in Denmark can contribute alleles to the European gene pool. A living collection of Phragmites genotypes was established in the experimental garden of Aarhus University beginning in The collection represents all continents and all species of the genus, but most samples are from Europe. The clones established in the garden are listed in \[ Supplementary Data \] of the present study. This collection was sampled for DNA and used for a phenological investigation. For the molecular work we included additional reference samples not cultivated in the garden from the Mediterranean area and the Middle East from Sardinia-Italy, Libya and Jordan Lambertini et al. The distribution of the samples used in the study is shown in Fig. Previous studies Lambertini et al. Outgroups are necessary in any phylogenetic analysis to provide a point of comparison and polarize ancestral and derived characters. The inclusion of a single outgroup or a few distantly related ougroups may, however, not be helpful, as homologies and alignment might become uncertain. Therefore, we included a considerable number of outgroups which should increase the likelihood that the European basal node is correctly placed and that characters are correctly polarized. Distribution of the Phragmites samples used in the study. The sample set includes the species P. Seventy-three clones from the Aarhus and Mediterranean collections, selected to represent Europe from the Atlantic Coast to the Danube delta in longitude, and from northern Sweden to Jordan in latitude, were sequenced. The Danube delta, represented by several genotypes in our garden, was more densely sampled 20 samples because of the high genetic diversity detected in this region in previous studies Lambertini et al. The amplified product was sequenced with forward and reverse primers in an ABI sequencer. As most of the variation in these regions is represented by tandem repeats, we amplified and sequenced those samples differing from GenBank accessions multiple times, to make sure that repetitive sequences were not due to amplification or sequencing artefacts. Sequences were aligned with the program BioEdit ver. Saltonstall, pers. Variation in the number of mononucleotide repeats in the trn T- trn L sequence has previously been attributed to intra-haplotypic variation Saltonstall and, as such, has not been used to define haplotypes. However, given the frequency of these variants in the European data set, and the possible phylogeographic value of this variation Chu et al. The capital letter symbolizing the haplotypes e. The variation among these haplotypes is given by indels, substitutions and number of tandem repeat units of and bp motifs minisatellites. Haplotypes defined by a capital letter followed by a number e. M1, M2, M3, L1, O2, etc. These cp-microsatellite variants have not been deposited in GenBank because the classification system used in the present study is preliminary. The analysis of a larger cp-variation in microstatellites worldwide might produce a different classification system K. Saltonstall and C. Lambertini, in preparation. In total, 69 different sequences were obtained combining our data set with the sequences downloaded from GenBank. To avoid overestimating the number of mutations associated with repeated sequences, we coded repeat motifs microsatellites and minisatellites as multistate characters Doyle In this way, differences due to the number of repeats are counted as one character, independently of their number. The final matrix counted characters, of which 27 were multistate characters. Amplified fragment length polymorphisms AFLPs and microsatellites or simple sequence repeat — SSRs data were obtained from previous global studies Lambertini et al. The amplified product was run on a 1. The three data sets were subject to parsimony analysis with PAUP ver. The matrix of genetic distances was imported in GenAlex ver. To increase the resolution of the European part of the PCoA, we plotted individual coordinate values in Excel graphs. Most of the Australian genotypes were excluded from the AFLP matrix of genetic distances because the PCoA failed to complete the analysis, as is the maximum number of samples that can be run by the program. However, as the Australian genotypes are closely related among each other, the placement of the main clusters on the PCoA is not affected by the exclusion of part of the genotypes of this group. A spectrum of genetic distances Rozenfeld et al. Samples from the native American cp-lineages and from the US Gulf Coast were excluded from the North American population, and samples from the Mediterranean region, Africa and Asia were excluded from the European group. In total, the North American population was represented by 30 samples, the European by and the Po Plain by Denmark is in the temperate area of the Northern Hemisphere. There are up to 18 daily hours of sunlight during the summer. Each genotype was planted in a separate pot of about 1 m diameter, and the pots were ordered in a square matrix at about 2 m distance from each other. All above-ground biomass was harvested and burnt at the end of the vegetative season to eliminate seed dispersal from the garden. All clones had been in the garden for at least 5 years before phenology was investigated. Given the weight of the pots, it was not possible to move them within the plot to control margin effect. However, given the large distance between pots, all genotypes had chances to receive pollen both from within the garden and from the wild. We recorded the reproductive condition of each genotype in and on a weekly basis from June to September. We were interested in knowing which genotypes of the garden, including European and non-European genotypes, could release pollen and seeds in Europe, and how dispersed or non-dispersed alleles contributed to the pattern of phylogeographic relationships within Europe and at the global scale. In November , we recorded whether or not genotypes were releasing seeds. Seed release was defined as the presence of depleted inflorescences visible bare inflorescence axes or easily detachable seeds upon touch. Cross- versus self-pollination was not considered relevant to the aim of the present study as alleles would have been present in the gene pool in both cases. AY , whereas the trn T- trn L sequences showed considerable variation. The limited variation in the rbc L-p sa I region was restricted to two base substitutions which distinguished haplotype AJ and haplotype AK. Conversely, the trn T- trn L sequences were constant within P. AY , while rbc L- psa I sequences detected two different haplotypes within the two P. European and Mediterranean haplotypes identified by cpDNA. The haplotypes are in alphabetical order. For example, trn T- trn L sequences 4a, 4c, 4d, etc. This variation in cp- microsatellites is also indicated in the number following the haplotype letter e. M1, M3, M4, etc. Based on the database of sequences accumulated from our previous and ongoing studies, many of the European haplotypes appeared to have intercontinental distribution, ranging from Africa to Europe, Far East Asia and North America. The most striking example is haplotype M, which is present on all four continents. Haplotype M presents several cp-microsatellite variants that are mainly distributed in the Mediterranean region. One such variant, haplotype M1 or Delta-type Hauber et al. Haplotype AI, previously described as Greeny3-type Lambertini et al. Haplotype AD or Greeny2-type Lambertini et al. Haplotype O presents several cp-microsatellite variants in Far East Asia. Two of these variants occur in Northern Europe and the Mediterranean region. Two cp-microsatellite variants of haplotype L were found in Europe. Haplotype L1 appears widely spread across the continent Belgium, Finland and Romania , whereas haplotype L2 was found only in Sweden. HQ was found in Estonia and Romania, and one of its cp-microsatellite variants was found in Turkey. HQ only on Djerba island in Tunisia. Phragmites frutescens haplotypes are shared with the species P. Both haplotypes I and U are present in Asia, where they show cp-microsatellite variation. Interestingly, only one of the variants, here indicated as I2, is present in Greece P. Despite frequent long-distance dispersal and lack of support as a monophyletic clade Fig. The PCoA clearly separates four main clusters of haplotypes within the genus, which largely overlap in geographic distribution. Chloroplast DNA diversity in Phragmites worldwide. Cluster 2 includes specimens assigned to P. Cluster 3 includes specimens assigned to P. Distribution: Far East Asia. The distribution is limited to North America. The classification of haplotypes vfM2, vfM3, vfE1, vfE3 and vfE4 Vachon and Freeland ; Freeland and Vachon does not follow the same classification system as Saltonstall and followed in the present study. According to the present study, vfM2 corresponds to haplotype L1, vfM3 should have a new letter, vfE1 deposited in GenBank both as E1 and E2 corresponds to a cp-variant of haplotype AB Meadows and Saltonstall , haplotype vfE3 Vachon and Freeland ; Freeland and Vachon corresponds to a cp-microsatellite variant of haplotype E Saltonstall , whereas haplotype E4 Vachon and Freeland ; Freeland and Vachon should have a new letter. The distribution ranges are defined by combining our collection sites with Fig. The pattern of interrelationships revealed by nuclear markers mirrors the haplotypic structure detected by cpDNA sequences at the global scale, as the same groups are supported in parsimony analysis Figs 3 A and 4 A. The parsimony analysis of SSR data Fig. The geographic differentiation within this cluster of haplotypes I and U and the lack of support in some regions reveal that different pools of nuclear alleles evolved in different parts of the range. On the contrary, the cluster of European P. SSR alleles did not separate European P. Amplified fragment length polymorphism diversity in Phragmites worldwide. The clusters of genotypes are enumerated according to Fig. B and C focus on European P. A The different colours represent the different haplotypes in Fig. Chloroplast DNA cluster 1 Fig. Cluster 2 includes P. Cluster 3 includes P. B The different colours indicate different continents. North America is represented by two colours in order to distinguish between two different introductions from Eurasia haplotype M and from the Mediterranean region haplotype M1. Panel C focuses on the temperate region of Europe. The different colours indicate different haplotypes. SSR diversity in Phragmites worldwide. The clusters of genotypes are enumerated according to Figs 2 and 3 A. Cluster 1 includes hybrid genotypes of P. Cluster 4 P. B focuses on cluster 6. The different colours indicate different continents and different species. C focuses on the temperate region of Europe. Differently from the cpDNA sequences, nuclear alleles detected geographic structure within the European group. SSRs indicated relationships of Mediterranean P. A nuclear DNA band of about bp amplified by waxy primers appeared exclusive to the Mediterranean genotypes Fig. Phragmites frutescens , like P. Samples carrying both bands of and bp had previously been interpreted as hybrids between P. However, the sequence of the bp band isolated from this sample from Sardinia Italy did not match any of the five sequences of P. Diversity in grass - waxy sequences in the Mediterranean region and in Africa. The bp band distinguishes Mediterranean lanes 3—10 from European temperate P. The genotype in lane 2 is from Mallorca, Spain, and appears as one undifferentiated yellow genotype clustering together with temperate European and North American P. It also released seeds in Denmark. The bp band marks P. Samples with and bp bands lanes 9, 11 and 12 have been interpreted as hybrids between P. Senegal lanes 11—13 appears as a hybrid zone with hybrids and non-hybrids. Sardinia Italy lanes 9—10 also appears as a hybrid zone. The red group of Asiatic samples Fig. This group in North America includes genotypes from the Atlantic Coast. Of the genotypes present in the experimental area, only 83 reproduced in and and released seeds in \[see Supplementary Data \]. Genotypes of P. Within the European cluster cluster 3 , P. Two genotypes from Romania and three genotypes from the Mediterranean region more closely related to the temperate group of European P. Amplified fragment length polymorphism relationships among dispersing and non-dispersing genotypes in the common garden at Aarhus University. The red dots in the European gene pool belong to genotypes from South Africa, Senegal, the Mediterranean region, North America introduced from the Mediterranean region and Romania. Compared with , the pattern of reproduction observed differed in November \[see Supplementary Data \]. Genotypes from Northern Europe, South Africa and Australia that did not produce inflorescences in , flowered in , and the Northern European and Australian genotypes released seeds in On the contrary, genotypes that produced seeds in produced only inflorescences in November genotypes from Northern Europe, Romania, Central Russia and Sakhalin Island, and one native North American genotype from Utah. The duration of the flowering period ranged from the end of August to mid-September for clones from Sakhalin Island, to flowering at the beginning of August and lasting until the end of September for most European genotypes. The longer flowering period of the European plants was the result of continuous inflorescence production and maturation among the ramets. In general, there was a shift in flowering times from north to south for the genotypes of the Northern Hemisphere. The few genotypes of the Southern Hemisphere that set seeds in Denmark started flowering at the end of August, and the first spikelets opened in early September Australia. Many late-flowering genotypes did not produce seeds in Denmark but they flowered and released pollen in and Chloroplast DNA sequences have identified four main clusters of haplotypes with continental or intercontinental distribution. This structure is also identifiable by nuclear markers. Haplotypes from two of these clusters are present in Europe and are sympatric in the Mediterranean region. In Europe, the two groups are recognized taxonomically, one being P. In total, 12 P. The two haplotypes indicated in the present study as vfM2 and vfM3, found by Vachon and Freeland in Great Britain and described in GenBank as M2 and M3 do not follow the alphabetical system introduced by Saltonstall According to the present study, vfM2 corresponds to haplotype L1, whereas vfM3 should have a new letter. Differently from Saltonstall , Vachon and Freeland and Saltonstall and Lambertini , we also considered cp-microsatellite variation in the analyses, because cp-microsatellite variation appeared restricted within limited regions in the range of some haplotypes, and may thus be phylogeographically informative. The presence of the same variant M1 also in Romania, as shown in the present study, is not surprising as the population in the US Gulf Coast shares several genetic similarities with the population in the Danube delta Lambertini et al. We would expect a random pattern of distribution and relationships if cp-microsatellite variation in haploytpe M were merely due to homoplasy. This, however, cannot be generalized to all haplotypes. The inclusion of cp-microsatellite variants in the analysis and the different codification system that we used for repeated sequences can explain differences with the global haplotypic structure previously presented by Saltonstall , Vachon and Freeland , Chu et al. This is the first study that compares cpDNA and nuclear variation at the global scale, and the fact that the major groups can be identified by both DNA sources indicates reliability of the molecular markers used and strongly supports the phylogeographic model presented in this study. Africa is differentiated from Europe. Differentiation is also ongoing in North America. The gap in our sampling from East Europe to the Far East does not allow us to set the phylogeographic limit of the European gene pool in Asia. However, Far East Phragmites appears different from European Phragmites both in haplotypic composition and nuclear markers. The study of Chu et al. In our data, haplotype P is represented in a separate cluster of haplotypes, which also has distinctive nuclear alleles cluster 2, Figs 2 , 3 A and B. More difficult to understand is the extension of the group to which P. Phragmites frutescens shares the cpDNA sequences with both P. This finding may have revealed an important role of P. However, many more genotypes within the cluster of haplotypes I and U cluster 1 need to be included in the analyses to understand the structure of this group. Within the temperate European region European P. Several closely related haplotypes co-exist in the area, but haplotypic structure is not identifiable by nuclear markers. This suggests long-distance dispersal of genotypes and extensive gene flow within the temperate region. The phenological results appear in agreement with the European phylogeographic pattern. The common garden study in Europe shows that there are limits to sexual reproduction and seed dispersal across Europe, and these limits appear to select the European genetic variation. Genotypes from the Southern Hemisphere can establish in Denmark, but only a few can occasionally conclude their reproductive cycle in Denmark. The genotypes from Australia started to flower late in Denmark September , but some of them released pollen and a few set seeds. Also one South African genotype started to flower in September but did not conclude its cycle. The fact that these geographically distant genotypes can occasionally reproduce in Denmark might explain some of the long-distance relationships seen in Europe, for example the phylogenetic relationship of haplotype K among South African, Spanish, Romanian and American genotypes. The European K-related genotypes have different nuclear alleles from the South African population, but not from the European gene pool. Another observation that suggests gene flow in Europe is given by the cytotypic variation within this cp-lineage. The South African individuals are octoploids whereas the Romanian genotypes are part of a mixed cytotype population in the Danube delta which includes octoploids Pauca-Comanescu et al. The same Romanian individuals analysed by flow cytometry by Clevering and Lissner were sequenced in the present study and cytotypic variation appeared distributed within every cp-lineage present in the region. Cytological incompatibilities might explain why the two Romanian genotypes do not reproduce in Denmark. Another possible explanation is that haplotype K was originally dispersed from Europe to South Africa where it eventually differentiated from the European pool. We cannot reconstruct the history of haplotype K based on this data set, but studies at the population level and experimental crosses can clarify gene flow possibilities and limitations. Given the north—south gradient in flowering times across Europe, reported also in other common garden studies Clevering et al. With the exception of one genotype from Gorgona Island Italy , which shared haplotype O with the North European genotypes and is likely to have been introduced from the mainland F. Masi, pers. This suggests either climatic barriers to dispersal of predominantly tropical and subtropical genotypes into the temperate regions of Europe, or breeding barriers that keep the Mediterranean genotypes isolated from temperate P. The nuclear waxy bands captured the diversity of the Mediterranean group identified by the bp waxy band , as well as its distinctiveness from European P. The homologies and relationships based on the DNA sequences contained in the waxy bands have proven phylogenetically informative between the African and American genotypes Lambertini et al. Several additional bands were amplified by grass-waxy primers in the Mediterranean samples, and these could provide further insights into the relationships of Mediterranean genotypes, as well as the DNA source of such bands. The occurrence of individuals with and bp bands, like the sample in lane no. SSR alleles detected similarities with both P. We could not find matching waxy sequences to conclusively demonstrate hybridization, but we also have a very small number of samples of these tropical species which span in distribution from South Africa, to Uganda, Burkina Faso, Thailand and Australia, and only two single individuals of P. More variation is likely to be present in the Mediterranean region. We recently found a herbarium accession of P. The specimen was collected in Misterbianco, close to Catania, in Sicily Italy in accession no. BOLO and has seeds. A focused study in the Mediterranean region is necessary to fully understand the relationships of Mediterranean P. The geographic distribution of the bp band matches the distribution of P. Both epithets appear therefore in agreement with the genetic identity of Mediterranean P. However, this needs to be confirmed with phenotypes of var. The two P. The introduced population of European Phragmites in North America also appears to be differentiated from Europe. This cluster of individuals, also detected by Lambertini et al. This study reveals European P. The large scale of the study opens several windows onto the genetic dynamics within the European group and the relationships within the genus that stimulate further research. The Mediterranean region appears an interesting spot of Phragmites diversity for evolutionary biologists, geneticists, taxonomists, ecologists and ecophysiologists. The Danube delta is another important region for its cytologic and haplotypic diversity, and could finally explain how chromosome numbers affect gene flow and diversification processes. Furthermore, given the genetic similarities of the Danube delta population to the population in the Mississippi delta Lambertini et al. The rest of Europe is no less interesting in the context of haplotypic diversity, morphology and long-distance relationships, and this should be taken into account in studies at the population scale. Phragmites australis has been farmed in Europe for thousands of years for roof thatching and more recently it is being extensively used in constructed wetlands. It would be particularly interesting to understand how human-assisted transport of genotypes has influenced the current pattern of phylogeographic relationships. At the global scale, there is a large amount of diversity that is missing from this phylogeographic model. Phragmites diversity in Africa, Asia and South America is under-represented in our study and in the literature. The literature about China and the Far East is increasing, but there is still a big geographic gap to be covered with samples, which will complete the genetic picture and provide more outgroups and hopefully statistical support to understand the relationships among the main Phragmites phylogeographic groups. Supplementary Data. Table 1. Phenological registrations of flowering times, seed setting and seed release in the common garden at Aarhus University, Denmark. Flowering times in and seed release in are reported. Every individual corresponds to a different genotype. Genotypes are divided in the main groups identified by the study and are listed from north to south within each group. All authors commented on and approved the manuscript. We thank Kristin Saltonstall and an anonymous reviewer for their personal engagement in the review of the manuscript and for the constructive inputs. Saltonstall is also thanked for classifying the new haplotypes. Professor P. Antioco Island, Sardinia, Italy. Laura A. Karin M. Kettenring is thanked for commenting on an early draft of the manuscript. Special thanks to all colleagues and friends that sent Phragmites leaves, seeds or rhizomes from different parts of the world and contributed to the creation of this sample set. Google Scholar. Google Preview. Oxford University Press is a department of the University of Oxford. It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide. Sign In or Create an Account. Sign in through your institution. Annals of Botany Company Journals. Advanced Search. Search Menu. Article Navigation. Close mobile search navigation Article Navigation. Volume Article Contents Abstract. Materials and methods. Conclusions and forward look. Additional information. Accession numbers. Sources of funding. Contributions by the authors. Conflict of interest statement. Journal Article. Exploring the borders of European Phragmites within a cosmopolitan genus. Department of Bioscience, Plant Biology. Oxford Academic. Brian K. Tenna Riis. Birgit Olesen. Hans Brix. Revision requested:. Select Format Select format. Permissions Icon Permissions. Abstract Background and aims. Open in new tab Download slide. Table 1 Open in new tab. Sample ref. NCBI accession no. Pa68AL P. Phylogeographic analyses of Phragmites australis in China: native distribution and habitat preference of the haplotype that invaded North America. Google Scholar Crossref. Search ADS. Growth of Lythrum salicaria and Phragmites australis plants originating from a wide geographical area: response to nutrient and water supply. Genetic diversity, ecophysiology and growth dynamics of reed Phragmites australis. Identification of natural hybrids in Korean Phragmites using haplotype and genotype analyses. Taxonomy, chromosome numbers, clonal diversity and population dynamics of Phragmites australis. 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Seasonal development of morphological and physiological characteristics at the natural site and after transplantation. A set of primers for the amplification of noncoding regions of chloroplast DNA in the grasses. Cryptic invasion by a non-native genotype of the common reed, Phragmites australis , into North America. Microsatellite variation within and among North American lineages of Phragmites australis. Remnant native Phragmites australis maintains genetic diversity despite multiple threats. Phragmites frutescens Gramineae re-visited. The discovery of an overlooked, woody grass in Greece, especially Crete. Universal primers for amplification of three non-coding regions of chloroplast DNA. Phylogeographic inferences from chloroplast DNA: quantifying the effects of mutations in repetitive and non-repetitive sequences. Issue Section:. Download all slides. Supplementary data. Supplementary Data - zip file. Comments 0. Add comment Close comment form modal. 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