Growth of Lythrum salicaria and Phragmites australis plants originating from a wide geographical area: response to nutrient and water supply

Aim The response of L. salicaria and P. australis plants originating from a broad latitudinal gradient to mineral nutrient and water supply was studied. We suggest implications for mechanisms possibly underlying the invasiveness of these two plant species. Location A common garden tub experiment was carried out at T?eboň, Czech Republic (49° N 14°47′ E, altitude 433 m). Methods Plants of 30 L. salicaria and 6 P. australis populations originating from a wide geographical area in Europe and in Israel (60°?32° N latitude, 6° W?20° E longitude) were cultivated for one growing season at two water levels and three (L. salicaria) or two (P. australis) nutrient doses, respectively. At the end of cultivation, basic morphological and growth characteristics were recorded. Results The latitude of the original geographical location was the most significant factor, affecting both the phenology and most of the plant morphological and growth characteristics measured in both plant species, with the characteristics related to plant size being negatively correlated with latitude. The effect of nutrient dose was very strong in both species, while the effect of water level was more pronounced in P. australis than in L. salicaria. Main conclusions The results confirmed the existence of a large phenotypic variability in both plant species within their native ranges of occurrence. In both plant species, the latitudinal variation in plant morphological and growth characteristics most probably indicates the differentiation of genotypes adapted to local geographical conditions. The plants of both species originating from all geographical locations tested responded to the respective nutrient treatments in a similar manner. Exceptions were found in growth characteristics related to reproduction in L. salicaria, indicating that better nutrient supply could enhance the reproductive ability of certain genotypes. Based on the results obtained, we suggest implications for mechanisms possibly underlying the invasiveness of the plant species studied.     

Comparison of quantitative and molecular genetic variation of native vs. invasive populations of purple loosestrife (Lythrum salicaria L., Lythraceae)

Study of adaptive evolutionary changes in populations of invasive species can be advanced through the joint application of quantitative and population genetic methods. Using purple loosestrife as a model system, we investigated the relative roles of natural selection, genetic drift and gene flow in the invasive process by contrasting phenotypical and neutral genetic differentiation among native European and invasive North American populations ( Q _ST −  F _ST analysis). Our results indicate that invasive and native populations harbour comparable levels of amplified fragment length polymorphism variation, a pattern consistent with multiple independent introductions from a diverse European gene pool. However, it was observed that the genetic variation reduced during subsequent invasion, perhaps by founder effects and genetic drift. Comparison of genetically based quantitative trait differentiation ( Q _ST) with its expectation under neutrality ( F _ST) revealed no evidence of disruptive selection ( Q _ST >  F _ST) or stabilizing selection ( Q _ST <  F _ST). One exception was found for only one trait (the number of stems) showing significant sign of stabilizing selection across all populations. This suggests that there are difficulties in distinguishing the effects of nonadaptive population processes and natural selection. Multiple introductions of purple loosestrife may have created a genetic mixture from diverse source populations and increased population genetic diversity, but its link to the adaptive differentiation of invasive North American populations needs further research.     

Impact and management of purple loosestrife (Lythrum salicaria) in North America

The invasion of non-indigenous plants is considered a primary threat to integrity and function of ecosystems. However, there is little quantitative or experimental evidence for ecosystem impacts of invasive species. Justifications for control are often based on potential, but not presently realized, recognized or quantified, negative impacts. Should lack of scientific certainty about impacts of non-indigenous species result in postponing measures to prevent degradation? Recently, management of purple loosestrife (Lythrum salicaria), has been criticized for (1) lack of evidence demonstrating negative impacts of L. salicaria, and (2) management using biocontrol for lack of evidence documenting the failure of conventional control methods. Although little quantitative evidence on negative impacts on native wetland biota and wetland function was available at the onset of the control program in 1985, recent work has demonstrated that the invasion of purple loosestrife into North American freshwater wetlands alters decomposition rates and nutrient cycling, leads to reductions in wetland plant diversity, reduces pollination and seed output of the native Lythrum alatum, and reduces habitat suitability for specialized wetland bird species such as black terns, least bitterns, pied-billed grebes, and marsh wrens. Conventional methods (physical, mechanical or chemical), have continuously failed to curb the spread of purple loosestrife or to provide satisfactory control. Although a number of generalist insect and bird species utilize purple loosestrife, wetland habitat specialists are excluded by encroachment of L. salicaria. We conclude that (1) negative ecosystem impacts of purple loosestrife in North America justify control of the species and that (2) detrimental effects of purple loosestrife on wetland systems and biota and the potential benefits of control outweigh potential risks associated with the introduction of biocontrol agents. Long-term experiments and monitoring programs that are in place will evaluate the impact of these insects on purple loosestrife, on wetland plant succession and other wetland biota.     

Plasticity of Lythrum salicaria and Phragmites australis growth characteristics across a European geographical gradient

Plants of Lythrum salicaria and Phragmites australis originating from localities across the European north–south geographical gradient were cultivated in parallel in an outdoor tub experiment. A strong correlation was found between growth and morphometric characteristics related to plant size (plant height, basal diameter, aboveground- and belowground plant biomass, etc.) and the position of the respective populations along the north–south gradient. Plants of both L. salicaria and P. australis from the southern localities grew taller and more vigorously and flowered later than plants from relatively more northern localities. From this point of view, the plants originating from south European populations were comparable to invasive North American plants. Our study indicates that explanation of the competitive success of populations invading new geographical areas may involve the role of geographic gradients within the species native range.     

Genetic differentiation in life-history traits of introduced and native common ragweed (Ambrosia artemisiifolia) populations

Introduced species represent opportunities to observe evolution over contemporary time scales, and as exotics encounter new environments, adaptive responses can occur, potentially contributing to invasion. Here, we compare 22 native North American populations and 12 introduced European populations of common ragweed (Ambrosia artemisiifolia) in five common gardens (control, herbivory, light stress, nutrient stress and drought). We found evidence for improved growth and reproduction of the introduced populations in most environments, particularly in the light stress. However, under drought conditions, the introduced plants experienced more rapid wilting and mortality than their native counterparts, evidence consistent with a life-history trade-off between rapid growth and drought tolerance. Moreover, we found parallel latitudinal clines in flowering time and correlations between fitness components and the local climate of the source populations in both ranges. Together these data provide evidence for adaptation to local environmental conditions in the native and introduced range of common ragweed.     

Latitudinal population differentiation in phenology,life history and flower morphology in the perennial herb Lythrum salicaria

Abstract In plants with a wide distribution, phenological characters can be expected to vary clinally along climatic gradients, whereas other characters important for adaptation to local biotic and abiotic factors may vary in a more mosaic fashion. We used common‐garden experiments and controlled crosses to examine population differentiation in phenology, life history and morphology in the perennial herb Lythrum salicaria along a latitudinal transect through Sweden (57°N to 66°N). Northern populations initiated growth and flowering earlier, flowered for a shorter period, were shorter, produced more and larger winter buds, and were older at first reproduction than southern populations. Flower morphology varied significantly among populations, but was, with the exception of calyx length, not significantly related to latitude of origin. Survival in the common garden (at 63°49′N) was positively correlated with latitude of origin and the size and number of winter buds produced in the preceding year. The results suggest that the among‐population differences in phenology and life history have evolved in response to latitudinal variation in length of the growing season. Further studies are required to determine whether population differentiation in flower morphology is maintained by selection.     

Isozyme characterization of genetic diversity in Minnesota populations of purple loosestrife,Lythrum salicaria (lythraceae)

Starch gel electrophoresis of plant proteins was used to identify purple loosestrife (Lythrum spp.) cultivars and weedy populations. Preliminary determinations were made as to what degree weedy loosestrife populations were related (or genetically similar) to populations of L. alatum, L. virgatum, and horticultural cultivars. Cluster analysis of the data indicated that native L. alatum was genetically different from all populations of purple loosestrife and cultivars examined. The L. salicaria and L. virgatum cultivars, as groups, were not genetically distinguishable from the weedy populations analyzed. Seven cultivars of L. salicaria origin analyzed as a group were not distinguishable from the eight cultivars of L. virgatum origin, indicating that separation by cultivar origin may not be feasible. While the two “groups” were not distinguishable, most individual cultivars could be distinguished from one another by isozyme phenotype. Genetic variation was high within populations of weedy purple loosestrife but low among populations, which is characteristic of polyploid, perennial plant species that are widely distributed. Geographic location did not consistently correlate with genetic similarity.     

Body size variation and caste ratios in geographically distinct populations of the invasive big‐headed ant,Pheidole megacephala (Hymenoptera: Formicidae)

Body size is an important life history trait that can evolve rapidly as a result of how species interact with each other and their environment. Invasive species often encounter vastly different ecological conditions throughout their introduced range that can influence relative investment in growth, reproduction and defence among populations. In this study, we quantified variation in worker size, morphology and proportion of majors among five populations of a worldwide invasive species, the big‐headed ant, Pheidole megacephala (Fabricius). The sampled populations differed in ant community composition, allowing us to examine if P. megacephala invests differently in the size and number of majors based on the local ant fauna. We also used genetic data to determine if these populations of P. megacephala represented cryptic species or if morphological differences could be attributed to change following introduction. We found significant variation in worker mass among the populations. Both major and minor workers were largest in Australia, where the ant fauna was most diverse, and minor workers were smallest in Hawaii and Mauritius, where P. megacephala interacted with few to no other ants. We also found differences in major and minor worker morphology among populations. Majors from Mauritius had significantly larger heads (width and length) relative to whole body size than those from Hawaii and Florida. Minors had longer heads and hind tibias in South Africa compared with populations from Australia, Hawaii and Florida. The proportion of majors did not differ among populations, suggesting that these populations may not be subject to trade‐offs in investment in major size versus number. Our molecular data place all samples within the same clade, supporting that these morphologically different populations represent the same species. These results suggest that the variation in shape and morphology of major and minor workers may therefore be the result of rapid adaptation or plastic responses to local conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 423–438.     

Biological control of the wetlands weed purple loosestrife (Lythrum salicaria) in the Pacific northwestern United States

Purple loosestrife (Lythrum salicaria) is an Eurasian perennial hydrophyte that has become naturalized in wetlands and in and along waterways throughout temperate North America. The ecological integrity of such areas is threatened by rapidly forming monotypic infestations that displace valued flora and diminish critical fish and wildlife habitat. The inability of physical, cultural, and chemical methods to provide adequate control of the weed has led to the development of an insect-based biological control program. The first field releases of the bud and leaf feeding beetles, Galerucella calmariensis and G. pusilla, and a root-mining weevil, Hylobius transversovittatus, were made in the United States and Canada in 1992. A total of 4740 Galerucella spp. adults were released in central Washington during 1992 and 1993 at eight sites and 471 H. transversovittatus egg inoculations were made in 1993 at three locations. Establishment of both Galerucella spp. was confirmed and Hylobius colonization was achieved.     

Genetic variation in flowering phenology and avoidance of seed predation in native populations of Ulex europaeus

The genetic variation in flowering phenology may be an important component of a species’ capacity to colonize new environments. In native populations of the invasive species Ulex europaeus, flowering phenology has been shown to be bimodal and related to seed predation. The aim of the present study was to determine if this bimodality has a genetic basis, and to investigate whether the polymorphism in flowering phenology is genetically linked to seed predation, pod production and growth patterns. We set up an experiment raising maternal families in a common garden. Based on mixed analyses of variance and correlations among maternal family means, we found genetic differences between the two main flowering types and confirmed that they reduced seed predation in two different ways: escape in time or predator satiation. We suggest that this polymorphism in strategy may facilitate maintain high genetic diversity for flowering phenology and related life‐history traits in native populations of this species, hence providing high evolutionary potential for these traits in invaded areas.     

The implications of accepting untestedhypotheses: a review of the effectsof purple loosestrife (Lythrum salicaria)in North America

The acceptance of poorly tested hypotheses has adverse scientific consequences, and may have adverse ecological and social consequences. The hypothesis that purple loosestrife (Lythrum salicaria) has deleterious effects on North American wetlands is an example. We traced the history of purple loosestrife and its control in North America and found little scientific evidence consistent with the hypothesis that purple loosestrife has deleterious effects. The most commonly cited study of the effects of purple loosestrife on native flora and fauna produced inconclusive results. The general acceptance of this hypothesis, however, has resulted in the introduction of nonindigenous insects for biological control. Efforts to control purple loosestrife may be misplayed and may have long-term ecological consequences if loosestrife does not have the impact it is believed to have. The acceptance of this hypothesis using scientific justifications may affect future scientific credibil ity. Careful evaluation of the precautionary principle is necessary when considering the control of nonindigenous organisms.     

Evidence for rapid evolution of phenology in an invasive grass

Evolutionary dynamics of integrative traits such as phenology are predicted to be critically important to range expansion and invasion success, yet there are few empirical examples of such phenomena. In this study, we used multiple common gardens to examine the evolutionary significance of latitudinal variation in phenology of a widespread invasive species, the Asian short‐day flowering annual grass Microstegium vimineum. In environmentally controlled growth chambers, we grew plants from seeds collected from multiple latitudes across the species' invasive range. Flowering time and biomass were both strongly correlated with the latitude of population origin such that populations collected from more northern latitudes flowered significantly earlier and at lower biomass than populations from southern locations. We suggest that this pattern may be the result of rapid adaptive evolution of phenology over a period of less than one hundred years and that such changes have likely promoted the northward range expansion of this species. We note that possible barriers to gene flow, including bottlenecks and inbreeding, have apparently not forestalled evolutionary processes for this plant. Furthermore, we hypothesize that evolution of phenology may be a widespread and potentially essential process during range expansion for many invasive plant species.     

Latitudinal variation in wild populations of Drosophila melanogaster: heritabilities and reaction norms

Large amounts of genetic variation for wing length and wing area were demonstrated both within and between Drosophila melanogaster populations along a latitudinal gradient in South America. Wing length and wing area showed a strong positive correlation with latitude in both wild flies and laboratory-raised descendants. Large population differences were observed for heritability and coefficient of variation of these two traits, whereas relatively small population differences were found for development time, viability, pupal mortality, sex ratio and their norms of reaction to four developmental temperatures. No clear-cut latitudinal clines were established for these life-history characters. These results are discussed in the light of Bergmann's Rule and the relation between larval development and adult body size.     

Evidence for association of chromosomal form and development time from complex clines and geographic races in the grasshopper Caledia captiva (Orthoptera: Acridadae)

Evidence for an association between chromosomal form and development time in the grasshopper Caledia captiva (F.) was obtained through comparison of two geographic taxa and analysis of a complex latitudinal cline within one of the taxa. Northern populations of the Moreton taxon possess a metacentric genome and are slow-developing. In contrast, the Torresian taxon, distributed throughout northern, coastal Australia, a region of pronounced seasonality in rainfall, and southern populations of the Moreton taxon, which inhabit a region of pronounced seasonality in temperature, Soth have an acrocentric genome and are fast-developing. The convergence of chromosomal form and development time between Torresian and southern Moreton populations appears to be driven by convergence in life history. Seasonality limits grasshoppers to one generation per year and favours fast development. The transition between relatively acrocentric southern Moreton populations and relatively metacentric northern Moreton populations is gradual but not monotonic. Instead, a shift to a bivoltine life history in the middle of the transect occurs and is associated with shifts in both development time and chromosomal form. These results imply an adaptive role for chromosomal form, although the causative link between chromosomal variation and variation in development time remains to be established.     

Habitat and Population Effects on the Germination and Early Survival of the Invasive Weed, Lythrum salicaria L. (purple loosestrife)

In this study, we examined how environmental factors and differences among populations of Lythrum salicaria L. (Lythraceae; purple loosestrife) influenced the establishment of this species into both its preferred wetland habitat and secondary upland habitat. Seeds and seedlings from five parents from each of the three populations were planted in each wetland and upland of two sites in both disturbed and undisturbed plots. More seeds germinated in wetland than in upland sites (31% as compared to 11%) and, although a disturbance resulted in greater germination (23% as compared to 17% in undisturbed plots), there was no interaction with habitats. The results of a simultaneous greenhouse germination study in which seeds were planted under saturated and drained conditions suggest that the higher germination observed in wetland sites was due to the higher moisture present at such sites (79% vs. 45% in saturated and drained conditions, respectively). Disturbance enhanced both germination and growth of L. salicaria and this was especially so for seedlings. Dry weights of seedlings from disturbed plots were five times greater than those from undisturbed plots ( =9.2 vs. 1.8g in disturbed and undisturbed conditions, respectively). In addition to environmental factors, population differences are likely to play an important role in the spread of L. salicaria as we found differences between the populations in both life stages examined. One of the populations germinated equally well in both moisture treatments. In our seedling establishment experiment, one of the populations produced both the smallest plants and the most and earliest flowers, suggesting superior reproduction.     

Body size variation among mainland populations of the western rattlesnake (Crotalus viridis)

In general, squamate reptiles follow the converse to Bergmann's rule, attaining smaller sizes in cooler environments, whereas other vertebrate groups follow Bergmann's rule, attaining larger sizes in cooler areas. Intensive studies of body size evolution for species of squamates are necessary to understand the processes responsible for this trend. Here I present data on body size variation among mainland populations of the western rattlesnake, Crotalus viridis. This species consists of two well-differentiated phylogenetic clades, therefore all analyses were performed for the C. viridis group as a whole and separately for each of the two clades within the C. viridis group. Although both phylogenetic and nonphylogenetic analyses were performed, the data did not show phylogenetic conservatism, and therefore the nonphylogenetic results are preferred. I found no significant relationships between mean adult female snout-vent length and any of the physical and climatic variables that were examined for the C. viridis group using simple linear regression analysis. Examined separately, I found that individuals of the western clade, C. oreganus, were smaller in cooler and more seasonal environments, whereas individuals of the eastern clade. C. viridis sensu stricto, were larger in cooler and more seasonal areas. Thus, the observed size trends were in opposite directions for the two clades. Multiple regression analysis revealed that seasonality was a stronger predictor of body size variation than was temperature for both clades. The differences in body size trends between these clades may be due to differences in mortality rates among populations.     

The evolution of trade-offs: geographic variation in call duration and flight ability in the sand cricket, Gryllus firmus

Quantitative genetic theory assumes that trade-offs are best represented by bivariate normal distributions. This theory predicts that selection will shift the trade-off function itself and not just move the mean trait values along a fixed trade-off line, as is generally assumed in optimality models. As a consequence, quantitative genetic theory predicts that the trade-off function will vary among populations in which at least one of the component traits itself varies. This prediction is tested using the trade-off between call duration and flight capability, as indexed by the mass of the dorsolateral flight muscles, in the macropterous morph of the sand cricket. We use four different populations of crickets that vary in the proportion of macropterous males (Lab = 33%, Florida = 29%, Bermuda = 72%, South Carolina = 80%). We find, as predicted, that there is significant variation in the intercept of the trade-off function but not the slope, supporting the hypothesis that trade-off functions are better represented as bivariate normal distributions rather than single lines. We also test the prediction from a quantitative genetical model of the evolution of wing dimorphism that the mean call duration of macropterous males will increase with the percentage of macropterous males in the population. This prediction is also supported. Finally, we estimate the probability of a macropterous male attracting a female, P, as a function of the relative time spent calling (P = time spent calling by macropterous male/(total time spent calling by both micropterous and macropterous male). We find that in the Lab and Florida populations the probability of a female selecting the macropterous male is equal to P, indicating that preference is due simply to relative call duration. But in the Bermuda and South Carolina populations the probability of a female selecting a macropterous male is less than P, indicating a preference for the micropterous male even after differences in call duration are accounted for.     

Effect of purple loosestrife (Lythrum salicaria) diet supplementation in rabbit nutrition on performance,digestibility, health and meat quality

In this study, 160 Hycole weaned rabbits (35 days old) were randomly divided into four groups of 40. The rabbits were studied throughout a 54-day experimentation period in order to determine the impact of dietary supplementation from herbs composed of 0.2%, 0.4% dry ground Lythrum salicaria leaves (LS) and 0.3% Cunirel® (CR; a commercial herb mixture containing LS as the main ingredient) on performance, digestibility, health and meat quality. The basal diet was given to the control group. No significant differences were found in performance, 10 rabbits from each group were selected for evaluation regarding apparent digestibility. The rabbits fed the control diet and the diet with the low level of LS had a higher level of CP digestibility than did the animals that were supplemented with the high LS levels and CR (85.7% and 84.9% v. 84.0% and 84.0%, respectively; P<0.05). The ether extract digestibility was lower in the treatment group with 0.4%LS addition and CR as compared with the control group (52.2% and 54.5% v. 62.6%, respectively; P<0.05). The slaughter process was performed on 89-day-old rabbits to study the carcass characteristics, meat quality, blood parameters, caecal contents and gut histology. The total leukocyte counts in the control animals were lower than they were in the rabbits fed 0.2%, 0.4%LS and CR (4.06 v. 8.25, 8.63 and 8.21×10⁹/l, respectively; P<0.05). For caecal fermentation, the caecal contents of the rabbits fed 0.4% of LS, showed higher concentrations of total volatile fatty acid (VFA; 24.1 v. 18.9 mg/kg dry matter (DM); P<0.05) and acetic acid (18.3 v. 14.4 mg/kg DM; P<0.05), but lower ammonia levels (594 v. 892 mg/kg DM; P<0.05) as compared with the control group. PCR-denaturing gradient gel electrophoresis analyses were performed to evaluate the microbial community in hard faeces, collected at days 35, 42, 49, 56, 70 and 89, whereas the caecal contents were taken after slaughtering. The results demonstrated that between the treatment groups, the similarity of the microbial communities was higher as compared with the control group. Moreover, only age was shown to influence microbiota diversity. In conclusion, the results of this study indicated that supplementation of LS in rabbit diets leads to an increase in the total white blood cells, total VFA and acetic acid concentration, and a decrease in the ammonia levels, as well as the digestibility when CR and high level of LS were supplemented, without causing any adverse effects on other parameters.     

Latitudinal variation of morphological characteristics in the swallowtail Sericinus montelus Gray, 1798 (Lepidoptera: Papilionidae)

Patterns of latitudinal variation in the phenotype or genotype of an organism may provide evidence for natural selection. In this study, we investigated seven populations of swallowtail Sericinus montelus Gray, 1798 (Lepidoptera: Papilionidae), a non‐migratory species, to explore the latitudinal variation of morphological characteristics in adults. The results showed that body size and the development of dark pigmentation on wings in this species responded strongly to latitude. The body size of both male and female adult of S. montelus was negatively correlated with latitude. These findings provided solid evidence to support the converse Bergmann's rule. We considered that the observed variation in morphological characteristics was most likely mediated by the seasonal length and thermoperiod to adapt to different latitudinal environment (e.g. shortened developmental time of immature stages for smaller body size at higher latitude). Moreover, the tendency towards progressively darker colour patterns (only in adult males) at increasingly low latitudes was consistent with Gloger's rule. We suggested that the observed colour variation was most likely associated with thermoregulation. Slight variation in the morphology of the W‐shaped stripe on the forewing of adult females was also found, and we presumed that the functions of sexual preferences, mimicry and thermoregulation might be involved.