Incidences of gross morphological disorders in barbel (Barbus barbus) in three rivers in southern England

Incidences of gross morphological disorders in barbel ( Barbus barbus ) were investigated in three rivers in southern England. The water quality at the sites chosen for study on the Rivers Teme, Kennet and Lee were classified (according to the criteria of the National Water Council, 1978) as 1A, IB and 2, respectively. Generally, low levels of gross morphological abnormalities were seen in fish caught in the Teme and Kennet, although there were incidences of ocular disorders and extra barbules in 10 and 7%, respectively, of the Kennet fish. No tumours were found in barbel from the River Teme whereas there was a 2% incidence in the Kennet fish. In contrast barbel from the River Lee showed high levels of all the gross morphological disorders, especially haemorrhagic patches on the ventral surface (36%) and ocular disorders (72%). Tumours were seen in 25% of the barbel caught in the River Lee. The incidences of most of these gross morphological disorders did not appear to be age related. The possibility that these disorders were related to water quality is discussed.     

Loss of genetic integrity correlates with stocking intensity in brook charr (Salvelinus fontinalis)

Supportive breeding and stocking performed with non‐native or domesticated fish to support sport fishery industry is a common practice throughout the world. Such practices are likely to modify the genetic integrity of natural populations depending on the extent of genetic differences between domesticated and wild fish and on the intensity of stocking. The purpose of this study is to assess the effects of variable stocking intensities on patterns of genetic diversity and population differentiation among nearly 2000 brook charr (Salvelinus fontinalis) from 24 lakes located in two wildlife reserves in Québec, Canada. Our results indicated that the level of genetic diversity was increased in more intensively stocked lakes, mainly due to the introduction of new alleles of domestic origin. As a consequence, the population genetic structure was strongly homogenized by intense stocking. Heavily stocked lakes presented higher admixture levels and lower levels of among lakes genetic differentiation than moderately and un‐stocked lakes. Moreover, the number of stocking events explained the observed pattern of population genetic structure as much as hydrographical connections among lakes in each reserve. We discuss the implications for the conservation of exploited fish populations and the management of stocking practices.     

Patterns of grassland invasions by trees: insights from demographic and genetic spatial analyses

Aims Woody invasions into grasslands have increased globally due to changing land use, climate and introduced woody species, but spatial processes generating and sustaining these invasions are not well understood. To gain insight into the patterns of spread of tree populations within grasslands, and to propose a full spatial analytical toolbox for studying native and non-native woody species spread when long-term data are not available, we tested if 50 years of grassland invasion in Western Carpathians by Norway spruce (Picea abies Karst.) proceeded by one of the two traditionally competing hypotheses of species spread: (i) by frontier expansion, or (ii) by advanced groups established ahead of the population frontier. We also tested whether the pattern of invasion changed over time.Methods We analyzed the spatial demographic and genetic patterns of a Norway spruce population invading a Western Carpathian grassland using Ripley's L (t) and genetic kinship coefficients (F ij). We mapped and genotyped spruce trees across the invasion front (from the invasion leading edge to fully colonized grassland near the source population) using three demographic classes (adults, juveniles and seedlings) to approximate the temporal aspects of the invasion. We studied how the spatial patterns of invasion by individual demographic classes and their genetic kinship varied among adjacent plots established at different distances from the source population (ranging from 0 to 160 m, in 40-m distance increments).Important findings Juveniles were positively genetically related to adults on fine scales (<4 m), suggesting that adults within the grassland acted as a seed source and accelerated early invasion. However, adults did not act as nucleation centers for the formation of advanced juvenile groups. Instead, genetically unrelated juveniles formed groups independently of adults. These groups were small and separate at the leading edge but they increased in size and graded into a continuous zone near the source population. Thus, juvenile recruitment occurred as a frontier expansion near the source population and as advanced groups controlled by environmental variation at the leading edge. Unlike juveniles, seedlings were clustered on all scales across the invasion front and formed groups around adult crowns at the invasion leading edge. The bulk of seedling establishment occurred at intermediate distances from the source population, independently from the adults, suggesting that the invasion front continued to expand as a frontier, gradually coalescing with the advanced groups at the leading edge. Thus, the grassland invasion was driven by a gradual frontier expansion of the original population during the first 50 years, with advanced groups enhancing but not driving the invasion process. Frontier expansion appeared more important as a mechanism of woody species spread early in the invasion process in this study, while advanced groups may play a larger role over longer temporal scales.     

Founding events in species invasions: genetic variation, adaptive evolution, and the role of multiple introductions

Invasive species are predicted to suffer from reductions in genetic diversity during founding events, reducing adaptive potential. Integrating evidence from two literature reviews and two case studies, we address the following questions: How much genetic diversity is lost in invasions? Do multiple introductions ameliorate this loss? Is there evidence for loss of diversity in quantitative traits? Do invaders that have experienced strong bottlenecks show adaptive evolution? How do multiple introductions influence adaptation on a landscape scale? We reviewed studies of 80 species of animals, plants, and fungi that quantified nuclear molecular diversity within introduced and source populations. Overall, there were significant losses of both allelic richness and heterozygosity in introduced populations, and large gains in diversity were rare. Evidence for multiple introductions was associated with increased diversity, and allelic variation appeared to increase over long timescales (~100 years), suggesting a role for gene flow in augmenting diversity over the long‐term. We then reviewed the literature on quantitative trait diversity and found that broad‐sense variation rarely declines in introductions, but direct comparisons of additive variance were lacking. Our studies of Hypericum canariense invasions illustrate how populations with diminished diversity may still evolve rapidly. Given the prevalence of genetic bottlenecks in successful invading populations and the potential for adaptive evolution in quantitative traits, we suggest that the disadvantages associated with founding events may have been overstated. However, our work on the successful invader Verbascum thapsus illustrates how multiple introductions may take time to commingle, instead persisting as a ‘mosaic of maladaptation’ where traits are not distributed in a pattern consistent with adaptation. We conclude that management limiting gene flow among introduced populations may reduce adaptive potential but is unlikely to prevent expansion or the evolution of novel invasive behaviour.     

Population genetic structure of the sidespot barb,Barbus neefi,from the north-eastern escarpment,South Africa

Allozyme analysis was used to determine patterns of genetic variation within and between populations of Barbus neefi. The products of 29 loci were analysed, with 17 loci being monomorphic in all populations. The genetic variability estimates compared well with values reported in the literature. The percentage of polymorphic loci (0.95 criterion) ranged from 0–20.69% and average expected heterozygosity from 0.004–0.069. Headstream populations generally revealed lower genetic variability than populations in low lying areas, which emphasizes the importance of conserving headstream populations. Fixed allele mobility differences were observed at the MPI-1 protein coding locus between the Selons River population and the other populations. The measures of genetic differentiation as assessed by F-statistics, the effective number of migrants per generation and Nei's unbiased genetic distance consistently separated the Selons River population from the rest. Nevertheless, these estimates all fell within the range considered for conspecific populations. There was also a clear genetic division between populations from north and south of the Olifants River. The three populations from the Dorps, Spekboom and Blyde Rivers (Limpopo River system) and the population from the Crocodile River (Incomati River system) showed negligible genetic differentiation. Human-assisted transfer of populations may be responsible for this close similarity.     

Contrasting patterns of genetic variation and structure in plant invasions of mountains

Aim To assess the population genetic consequences of the colonization of two species with contrasting mating systems, Solidago canadensis and Lactuca serriola, along altitudinal gradients in both their native and introduced ranges. Location Allegheny Mountains, West Virginia and Wallowa Mountains, Oregon, USA; Valais, southern Switzerland. Methods Leaf material was collected from populations along altitudinal gradients and genotyped at seven microsatellite loci for each species. Differences in variability between native and introduced areas and in relation to altitude were analysed using linear models. Differences in the genetic, geographical and altitudinal structure of populations between areas were analysed by AMOVA, cluster analysis and Mantel tests. Results Genetic variation within and across populations of S. canadensis was significantly reduced, while populations of L. serriola were significantly more variable, in the introduced area. Genetic diversity decreased significantly with altitude for S. canadensis but not L. serriola. Genetic structure of S. canadensis was similar in both areas, and populations were isolated by geographical but not altitudinal distance. By contrast, population structure of L. serriola was much weaker in the introduced area, and populations were not isolated by distance in either area. Main conclusions Solidago canadensis has experienced a strong genetic bottleneck on introduction to the Valais, but this has not prevented it from colonizing a wide altitudinal range. Variation in neutral markers is therefore not necessarily a good measure for judging the ecological behaviour of a species. By contrast, the greater variability of L. serriola in the introduced area, where it also occurs over a greater altitudinal range, can be explained by increased outcrossing among admixed populations. This suggests that the ecological amplitude of alien species might be enhanced after population admixture in the new range, especially for species with highly structured native populations. However, even genetically depauperate introduced populations can be expected to colonize the same environmental range that they occupy in the native area.     

Unintentional selection,unanticipated insights: introductions,stocking and the evolutionary ecology of fishes

Natural environmental change has produced countless opportunities for species to disperse into and persist in habitats where they previously did not exist. Introduction and stocking programmes have facilitated similar sorts of colonization opportunities across considerably greater geographical scales and often in much shorter periods of time. Even though the mechanism of colonization differs, the result can be the same: evolutionary change in the colonizing population in response to novel selection pressures. As a consequence, some human‐mediated fish transfers have unintentionally yielded novel research opportunities to study how phenotypes and genes interact with their environment and affect ecological and evolutionary change. The primary purpose here is to explore how work, directly or indirectly involved with human‐mediated transfers, has unintentionally yielded novel research and research opportunities in fish ecology and evolution. Insights have produced new knowledge or altered previously held perceptions on topics such as local adaptation, rate of evolutionary change, phenotypic plasticity, alternative reproductive strategies, population structure and colonization probability. Well‐documented stocking programmes, especially in terms of history, numbers and original population sources, can provide highly fertile ground for generating further insights on the ecology and evolution of fishes and of the factors likely to influence the success of conservation‐based, restoration programmes.     

Population genetics of an invasive species,Heracleum mantegazzianum: implications for the role of life history,demographics and independent introductions

The population genetic structure of an invasive, nonindigenous riparian weed (Heracleum mantegazzianum) in the northeast of England was investigated using microsatellite DNA markers. Data were used to assess the size and frequency of introductions into three catchments and the subsequent spread. We sampled 13 populations, including a remote population (Perivale, London) for comparison. Five loci were screened and considerable variation was found. Results revealed greater overall variation between populations from different catchments than those in the same catchment, and suggested the generation of population structure in the relatively short time since the initial introduction. Between-catchment variation may reflect population structure generated by local founders as the species spread and may indicate a large initial founder population at the time of the introduction into Britain, or multiple introductions. Within-catchment variation was consistent with expectations based on the water-borne dispersal of seeds in this species, and the relatively small dispersal range of likely pollinators. Independent introductions stand out in some cases as exceptions to the general pattern. Taken together the results are consistent with a relatively large initial founder population, and the subsequent spread of the species in local founder populations, followed by some level of inbreeding within local populations and novel introductions in some localities.     

Ecological and genetic differentiation of Barbus callensis populations in Tunisia

Enzyme electrophoresis on horizontal starch gel was carried out on 356 barbel individuals. The sampling comprised 278 individuals of the species Barbus callensis from 10 rivers in Tunisia. The other individuals belonged to reference species (outgroups) from France and Morocco. An ecological study was also carried out on Tunisian rivers. The results show a clear differentiation of the two samples from northwestern Tunisia, which was only partly correlated with ecological characteristics of the rivers they inhabit. There is no genetic cline, but rather a discontinuity between populations in the northwestern-most watershed and all the other Tunisian populations. This differentiation probably has a paleohistoric origin not only related to adaptation to ecological conditions but also to difficulties in colonizing the watersheds. The results do not indicate clearly a colonization direction for the genus Barbus in North Africa. Analysis of the Algerian populations would appear to be indispensable. Lastly, in contrast with the usual taxonomy, Morocco and Tunisia are populated by two closely related species, but B. callensis should remain the name of the Tunisian species, which was the first to be described in the small El Kebir basin, a river that flows from Tunisia to Algeria.     

Low genetic differentiation among morphologically distinct Labeobarbus species (Teleostei: Cyprinidae) in the Lake Victoria and Albertine basins,Uganda: insights from mitochondrial DNA

Labeobarbus altianalis and L. bynni bynni are hexaploid cyprinid fishes in the genus Labeobarbus. In the Great Lakes region of Africa, these two large-bodied barbs exhibit considerable morphological variations. Their intraspecific classification, currently based on geographical distribution and morphological variation, is of limited application due to the overlapping nature of the water systems. Patterns of genetic variation in the Albertine and Victoria drainage basins in Uganda were assessed using mitochondrial sequences of the control region (470 basepairs) and the cytochrome b gene (627 basepairs). At the control region 36 distinct haplotypes were observed in a total of 177 individuals. Relatively low overall nucleotide diversity (π = 0.010) was observed. Hierarchical analysis of the molecular variance revealed significant differentiation between populations in the total sample (F _ST = 0.172, p < 0.001), among populations within each basin (F _SC = 0.047, p < 0.001), but no significant subdivision among basins (F _CT = 0.131, p > 0.05). Phylogenetic analysis of the control region haplotypes resulted in a star-shaped phylogeny, with no clusters that reflect current taxonomic designation, morphotypes or basins. Phylogenetic analysis of cytochrome b gene sequences resulted in a tree topology similar to that observed for the control region. Thus the morphological characteristics separating the two species are intraspecific variation and the whole sample probably belongs to the same species.     

Diversity and distribution of genetic variation in gammarids: Comparing patterns between invasive and non‐invasive species

Biological invasions are worldwide phenomena that have reached alarming levels among aquatic species. There are key challenges to understand the factors behind invasion propensity of non‐native populations in invasion biology. Interestingly, interpretations cannot be expanded to higher taxonomic levels due to the fact that in the same genus, there are species that are notorious invaders and those that never spread outside their native range. Such variation in invasion propensity offers the possibility to explore, at fine‐scale taxonomic level, the existence of specific characteristics that might predict the variability in invasion success. In this work, we explored this possibility from a molecular perspective. The objective was to provide a better understanding of the genetic diversity distribution in the native range of species that exhibit contrasting invasive propensities. For this purpose, we used a total of 784 sequences of the cytochrome c oxidase subunit I of mitochondrial DNA (mtDNA‐COI) collected from seven Gammaroidea, a superfamily of Amphipoda that includes species that are both successful invaders (Gammarus tigrinus, Pontogammarus maeoticus, and Obesogammarus crassus) and strictly restricted to their native regions (Gammarus locusta, Gammarus salinus, Gammarus zaddachi, and Gammarus oceanicus). Despite that genetic diversity did not differ between invasive and non‐invasive species, we observed that populations of non‐invasive species showed a higher degree of genetic differentiation. Furthermore, we found that both geographic and evolutionary distances might explain genetic differentiation in both non‐native and native ranges. This suggests that the lack of population genetic structure may facilitate the distribution of mutations that despite arising in the native range may be beneficial in invasive ranges. The fact that evolutionary distances explained genetic differentiation more often than geographic distances points toward that deep lineage divergence holds an important role in the distribution of neutral genetic diversity.     

Natural hybridization of two species of tetraploid barbels: Barbus meridionalis and Barbus barbus (Osteichtyes, Cyprinidae) in southern France

Natural hybridization of Barbus barbus and B. meridionalis has been demonstrated in southern France. A genetical study (isozyme electrophoresis) of these species and their hybrids revealed the characteristics of hybridization. Ten enzyme markers and five morphological parameters were used to distinguish between the two parent species. Enzymatic divergence between the parent species included fixed allelic differences at certain loci and reduction in enzymatic activity, including the silencing of certain genes. A morphological study revealed a good correlation between the isozyme markers and morphological characters. Backcrosses were observed and this raises the question of the integrity of the parent species.     

Invasion genetics of the Eurasian spiny waterflea: evidence for bottlenecks and gene flow using microsatellites

The Eurasian spiny waterflea (Bythotrephes longimanus) is a predacious zooplankter that has increased its range in Europe and is rapidly invading inland water-bodies throughout North America's Great Lakes region. To examine the genetics of these invasions, we isolated five microsatellite DNA loci with between 5 and 19 alleles per locus. We sampled three populations where B. longimanus has been historically present (Switzerland, Italy, and Finland) as well as an introduced European population (the Netherlands) and three North American populations (Lakes Erie, Superior, Shebandowan). Consistent with a bottleneck during colonization (i.e. founder effect), average heterozygosities of the four European populations ranged from 0.310 to 0.599, and were higher than that of three North American populations (0.151-0.220). Pairwise F(ST) estimates among North American populations (0.002-0.063) were not significantly different from zero and were much lower than among European populations (0.208-0.474). This is consistent with a scenario of high gene flow among North American populations relative to that of European ones. Contrary to an invasion bottleneck, however, Erie and Superior populations contained similar numbers of rare alleles as European populations. Assignment tests identified several migrant genotypes in all introduced populations (the Netherlands, Erie, Superior, Shebandowan), but rarely in native ones (Switzerland, Italy and Finland). A large number of genotypes from North America were assigned to our Italian population suggesting a second, previously unidentified, invasion source somewhere in the region of northern Italy. Together, our results support an invasion bottleneck for North American populations that has been largely offset by gene flow from multiple native sources, as well as gene flow among introduced populations.     

Strong intraspecific variation in genetic diversity and genetic differentiation in Daphnia magna: the effects of population turnover and population size

Theory predicts that genetic diversity and genetic differentiation may strongly vary among populations of the same species depending on population turnover and local population sizes. Yet, despite the importance of these predictions for evolutionary and conservation issues, empirical studies comparing high‐turnover and low‐turnover populations of the same species are scarce. In this study, we used Daphnia magna, a freshwater crustacean, as a model organism for such a comparison. In the southern/central part of its range, D. magna inhabits medium‐sized, stable ponds, whereas in the north, it occurs in small rock pools with strong population turnover. We found that these northern populations have a significantly lower genetic diversity and higher genetic differentiation compared to the southern/central populations. Total genetic diversity across populations was only about half and average within‐population diversity only about a third of that in southern/central populations. Moreover, an average southern population contains more genetic diversity than the whole metapopulation system in the north. We based our analyses both on silent sites and microsatellites. The similarity of our results despite the contrasting mutation rates of these markers suggests that the differences are caused by contemporary rather than by historical processes. Our findings show that variation in population turnover and population size may have a major impact on the genetic diversity and differentiation of populations, and hence may lead to differences in evolutionary processes like local adaptation, hybrid vigour and breeding system evolution in different parts of a species range.     

Understanding trends in biological invasions by introduced mammals in southern South America: a review of research and management

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Patterns of genetic diversity reveal multiple introductions and recurrent founder effects during range expansion in invasive populations of Geranium carolinianum (Geraniaceae)

Genetic diversity, and thus the adaptive potential of invasive populations, is largely based on three factors: patterns of genetic diversity in the species'' native range, the number and location of introductions and the number of founding individuals per introduction. Specifically, reductions in genetic diversity (‘founder effects'') should be stronger for species with low within-population diversity in their native range and few introductions of few individuals to the invasive range. We test these predictions with Geranium carolinianum, a winter annual herb native to North America and invasive in China. We measure the extent of founder effects using allozymes and microsatellites, and ask whether this is consistent with its colonization history and patterns of diversity in the native range. In the native range, genetic diversity is higher and structure is lower than expected based on life history traits. In China, our results provide evidence for multiple introductions near Nanjing, Jiangsu province, with subsequent range expansion to the west and south. Patterns of genetic diversity across China reveal weak founder effects that are driven largely by low-diversity populations at the expansion front, away from the introduction location. This suggests that reduced diversity in China has resulted from successive founder events during range expansion, and that the loss of genetic diversity in the Nanjing area was mitigated by multiple introductions from diverse source populations. This has implications for the future of G. carolinianum in China, as continued gene flow among populations should eventually increase genetic diversity within the more recently founded populations.     

Contrasting patterns in genetic diversity following multiple invasions of fresh and brackish waters

Biological invasions may combine the genetic effects of population bottlenecks and selection and thus provide valuable insight into the role of such processes during novel environmental colonizations. However, these processes are also influenced by multiple invasions, the number of individuals introduced and the degree of similarity between source and receiving habitats. The amphipod Gammarus tigrinus provides a useful model to assess these factors, as its invasion history has involved major environmental transitions. This species is native to the northwest Atlantic Ocean, although it invaded both brackish and freshwater habitats in the British Isles after introduction more than 65 years ago. It has also spread to similar habitats in Western Europe and, most recently, to Eastern Europe, the Baltic Sea, and the Laurentian Great Lakes. To examine sources of invasion and patterns of genetic change, we sampled populations from 13 native estuaries and 19 invaded sites and sequenced 542 bp of the mitochondrial COI gene. Strong native phylogeographical structure allowed us to unambiguously identify three allopatrically evolved clades (2.3-3.1% divergent) in invading populations, indicative of multiple introductions. The most divergent clades occurred in the British Isles and mainland Europe and were sourced from the St Lawrence and Chesapeake/Delaware Bay estuaries. A third clade was found in the Great Lakes and sourced to the Hudson River estuary. Despite extensive sampling, G. tigrinus did not occur in freshwater at putative source sites. Some European populations showed reduced genetic diversity consistent with bottlenecks, although selection effects cannot be excluded. The habitat distribution of clades in Europe was congruent with the known invasion history of secondary spread from the British Isles. Differences in salinity tolerance among lineages were suggested by patterns of habitat colonization by different native COI clades. Populations consisting of admixtures of the two invading clades were found principally at recently invaded fresh and brackish water sites in Eastern Europe, and were characterized by higher genetic diversity than putative source populations. Further studies are required to determine if these represent novel genotypes. Our results confirm that biological invasions need not result in diminished genetic diversity, particularly if multiple source populations, each with distinctive genetic composition, contribute to the founding populations.