Morphological variability of black bullhead Ameiurus melas in four non‐native European populations

External morphology in black bullhead Ameiurus melas, a fish species considered to have high invasive potential, was studied in its four non‐native European populations (British, French, Italian and Slovak). The aim of this study was to examine this species' variability in external morphology, including ontogenetic context, and to evaluate its invasive potential. Specimens from all non‐native populations reached smaller body size compared to individuals from native populations. Juvenile A. melas were found to have a relatively uniform body shape regardless of the population's origin, whereas adults developed different phenotypes depending upon location. Specimens from the U.K., Slovak and French populations appeared to be rather similar to each other, whereas the Italian population showed the most distant phenotype. This probably results from the different thermal regime in the Italian habitat. Ameiurus melas from non‐native European populations examined in this study showed some potential to alter the body shape both within and between populations. The phenotypic plasticity of A. melas, however, was not found to be as significant as in other invasive fish species. The results suggest that morphological variability itself is not necessarily essential for invasive success. The invasiveness of A. melas is therefore probably favoured by variations in its life‐history traits and reproduction variables, together with some behavioural traits (e.g. voracious feeding and parental care) rather than by phenotypic plasticity expressed in external morphology.     

Plasticity and genotypic variation in some Mentha × verticillata hybrids

Leaf and flower oil terpene composition and several plant morphological characteristics of 17 Mentha × verticillata hybrids were analysed during two growing seasons (1988 and 1989). The data obtained were used to study the phenotypic plasticity, the genotypic variation and the genetic variation for phenotypic plasticity. All plants showed high leveis of phenotypic plasticity for both oil chemical and morphometrical parameters. Higher degrees of genotypic variation were found among the plants for oll components while a higher phenotypic plasticity was observed for morphological parameters. Temperatures and rainfall data were collected during the growing seasons and correlated to the data obtained from plant oil and morphology. Low levels of phenotypic plasticity and high degrees of genotypic variation were found to form outliers in the population of M. x verticillata hybrids. The results obtained confirm a significant effect of environmental conditions on the physiology and morphology of the genus Mentha.     

A review of growth and life-history traits of native and non-native European populations of black bullhead <Emphasis Type="Italic">Ameiurus melas</Emphasis>

North American black bullhead, Ameiurus melas, which were introduced to Europe in the nineteenth and twentieth centuries, have received relatively little study. With focus on growth and reproduction, this extensive review, which includes new European data, aims to inform the risk analysis process concerning this non-native species in Europe. Surprisingly, the new data for Europe were more comprehensive than for native populations, with data available mainly from Oklahoma, and North and South Dakota (USA). In terms of relative growth, juvenile A. melas were found to have a relatively uniform body shape regardless of the population’s origin, whereas adults developed different phenotypes depending upon location. Overall growth trajectory was significantly faster for native than for non-native populations. Growth index values decreased significantly with increasing latitude in non-native but not native populations—the latter decreasing weakly with increasing altitude in the populations located at latitudes <40°. Mean general condition (slope ‘b’), mean sex ratio and mean egg diameter did not differ significantly between native and non-native populations. Absolute fecundity was slightly (but not significantly) higher in non-native than native populations. GSI data, which were very scarce for native populations, suggest gonad production may be slightly higher in native than in non-native populations. Precise data on age at maturity (AaM) are lacking for the native range, where 2–5 years is reported. Whereas, in the introduced range the greatest AaM was 3.5 years, and AaM decreases with increasing juvenile growth (TL at age 3). The populations with fastest juvenile growth tended to be from warmer water bodies where they are considered to be invasive. The great growth and life-history plasticity of black bullhead affords the species great potential to invade and establish viable populations in new areas.     

Rapid evolution in response to introduced predators II: the contribution of adaptive plasticity

Background  

Introductions of non-native species can significantly alter the selective environment for populations of native species, which can respond through phenotypic plasticity or genetic adaptation. We examined phenotypic and genetic responses of Daphnia populations to recent introductions of non-native fish to assess the relative roles of phenotypic plasticity versus genetic change in causing the observed patterns. The Daphnia community in alpine lakes throughout the Sierra Nevada of California (USA) is ideally suited for investigation of rapid adaptive evolution because there are multiple lakes with and without introduced fish predators. We conducted common-garden experiments involving presence or absence of chemical cues produced by fish and measured morphological and life-history traits in Daphnia melanica populations collected from lakes with contrasting fish stocking histories. The experiment allowed us to assess the degree of population differentiation due to fish predation and examine the contribution of adaptive plasticity in the response to predator introduction.     

Genotype–environment interaction and the ontogeny of diet-induced phenotypic plasticity in size and shape of Melanoplus femurrubrum (Orthoptera: Acrididae)

The developmental origin of phenotypic plasticity in morphological shape can be attributed to environment-specific changes in growth of overall body size, localized growth of a morphological structure or a combination of both. I monitored morphological development in the first four nymphal instars of grasshoppers (Melanoplus femurrubrum) raised on two different plant diets to determine the ontogenetic origins of diet-induced phenotypic plasticity and to quantify genetic variation for phenotypic plasticity. I measured diet-induced phenotypic plasticity in body size (tibia length), head size (articular width and mandible depth) and head shape (residual articular width and residual mandible depth) for grasshoppers from 37 full-sib families raised on either a hard plant diet (Lolium perenne) or a soft plant diet (Trifolium repens). By the second to third nymphal instar, grasshoppers raised on a hard plant diet had significantly smaller mean tibia length and greater mean residual articular width (distance between mandibles adjusted for body size) compared with full-sibs raised on a soft plant diet. However, there was no significant phenotypic plasticity in mean unadjusted articular width and mandible depth, and in mean residual mandible depth. At the population level, development of diet-induced phenotypic plasticity in grasshopper head shape is mediated by plastic changes in allocation to tissue growth that maintain growth of head size on hard, low-nutrient diets while reducing growth of body size. Within the population, there was substantial variation in the plasticity of growth trajectories since different full-sib families developed phenotypic plasticity of residual articular width through different combinations of head and body size growth. Genetic variation for diet-induced phenotypic plasticity of residual articular width, residual mandible depth and tibia length, as estimated by genotype–environment interaction, exhibited significant fluctuation through ontogeny (repeated measures MANOVA , family × plant × instar, P < 0.01). For example, there was significant genetic variation for phenotypic plasticity of residual articular width in the third nymphal instar, but not earlier or later in ontogeny. The observed patterns of genetic variation are discussed with reference to short-term constraints and the evolution of phenotypic plasticity.     

Phenotypic plasticity and genetic isolation-by-distance in the freshwater mussel <Emphasis Type="Italic">Unio pictorum</Emphasis> (Mollusca: Unionoida)

Freshwater mussels (Unionoida) show high intraspecific morphological variability, and some shell morphological traits are believed to be associated with habitat conditions. It is not known whether and which of these ecophenotypic differences reflect underlying genetic differentiation or are the result of phenotypic plasticity. Using 103 amplified fragment length polymorphism (AFLP) markers, we studied population genetics of three paired Unio pictorum populations sampled from two different habitat types (marina and river) along the River Thames. We found genetic differences along the Thames which were consistent with a pattern of isolation by distance and probably reflect limited dispersal via host fish species upon which unionoid larvae are obligate parasites. No consistent genetic differences were found between the two different habitat types suggesting that morphological differences in the degree of shell elongation and the shape of dorso-posterior margin are caused by phenotypic plasticity. Our study provides the first good evidence for phenotypic plasticity of shell shape in a European unionoid and illustrates the need to include genetic data in order properly to interpret geographic patterns of morphological variation.     

AFLP analysis shows high incongruence between genetic differentiation and morphology‐based taxonomy in a widely distributed tortoise

Morphology has traditionally been used to diagnose the taxa of various taxonomic ranks. However, there is growing evidence that morphology is not always able to reveal cryptic taxa, and that pronounced morphological variation could reflect phenotypic plasticity rather than evolutionary divergence. Spur‐thighed tortoises (the Testudo graeca complex), distributed in the western Palaearctic region, are characterized by high morphological variability and complicated taxonomy, which are under debate. Previous molecular studies using mainly mitochondrial DNA (mtDNA) sequences have revealed incongruence between genetic differentiation and morphology‐based taxonomy, suggesting that morphological variability is the result of phenotypic plasticity and stabilizing selection, which masks the true genealogies. In the present study, we used a range‐wide sampling and nuclear Amplified fragment length polymorphism (AFLP) markers to investigate genetic differentiation within the T. graeca complex. We found that spur‐thighed tortoises are differentiated into four geographically well‐defined AFLP groups: Balkans–Middle Eastern, western Mediterranean, Caucasian and central‐eastern Iranian. Compared with the distribution of mtDNA lineages, the groups are largely concordant, although the AFLP markers are less sensitive and distinguish fewer groups than do mtDNA sequences. The AFLP groups show an allopatric or parapatric distribution. The AFLP differentiation conflicts with the previously proposed morphology‐based taxonomy of the complex, suggesting that local adaptation to different environmental conditions may have led to the great extent of morphological variation within the same lineages. We propose a re‐evaluation of the taxa that were confirmed genetically using a thorough morphological analysis corrected for phenotypic plasticity. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Costs and limits of phenotypic plasticity: Tests with predator-induced morphology and life history in a freshwater snail

Potential constraints on the evolution of phenotypic plasticity were tested using data from a previous study on predator-induced morphology and life history in the freshwater snail Physa heterostropha. The benefit of plasticity can be reduced if facultative development is associated with energetic costs, developmental instability, or an impaired developmental range. I examined plasticity in two traits for 29 families of P. heterostropha to see if it was associated with growth rate or fecundity, within-family phenotypic variance, or the potential to produce extreme phenotypes. Support was found for only one of the potential constraints. There was a strong negative selection gradient for growth rate associated with plasticity in shell shape (β = ?0.3, P < 0.0001). This result was attributed to a genetic correlation between morphological plasticity and an antipredator behavior that restricts feeding. Thus, reduced growth associated with morphological plasticity may have had unmeasured fitness benefits. The growth reduction, therefore, is equivocal as a cost of plasticity. Using different fitness components (e.g., survival, fecundity, growth) to seek constraints on plasticity will yield different results in selection gradient analyses. Procedural and conceptual issues related to tests for costs and limits of plasticity are discussed, such as whether constraints on plasticity will be evolutionarily ephemeral and difficult to detect in nature.     

Genetic and morphological divergence at different spatiotemporal scales in the grasshopper Mioscirtus wagneri (Orthoptera: Acrididae)

The study of the association between morphological and genetic divergence can provide important information on the factors determining population differentiation and gene flow at different spatiotemporal scales. In this study we analyze the congruence between morphological and genetic divergence in the Iberian populations of Mioscirtus wagneri, a specialized grasshopper exclusively inhabiting highly fragmented hypersaline low grounds. We have found strong morphological variation among the studied localities and among mtDNA- and microsatellite-based genetic clusters. However, we have detected some cases of morphological convergence between highly differentiated populations. By contrast, certain genetically homogeneous populations at both mtDNA and microsatellite markers showed significant morphological differentiation which may be explained by phenotypic plasticity or divergent selection pressures acting at different spatiotemporal scales. Mantel tests also revealed that morphological divergence was associated with microsatellite- but not with mtDNA-based genetic distances. Overall, this study suggests that morphological traits can provide additional information on the underlying population genetic structure when only data on scarcely variable mtDNA markers is available. Thus, morphology can retain useful information on genetic structure and has the benefit over molecular methods of being inexpensive, offering a preliminary/complementary useful criterion for the establishment of management units necessary to guide conservation policies.     

Morphological anti‐predator defences in the nine‐spined stickleback: constitutive,induced or both?

Environmental differences among populations are expected to lead to local adaptation, while spatial or temporal environmental variation within a population will favour evolution of phenotypic plasticity. As plasticity itself can be under selection, locally adapted populations can vary in levels of plasticity. Nine‐spined stickleback (Pungitius pungitius) originating from isolated ponds (low piscine predation risk, high competition) vs. lake and marine populations (high piscine predation risk, low competition) are known to be morphologically adapted to their respective environments. However, nothing is known about their ability to express phenotypic plasticity in morphology in response to perceived predation risk or food availability/competition. We studied predator‐induced phenotypic plasticity in body shape and armour of marine and pond nine‐spined stickleback in a factorial common garden experiment with two predator treatments (present vs. absent) and two feeding regimes (low vs. high). The predation treatment did not induce any morphological shifts in fish from either habitat or food regime. However, strong habitat‐dependent differences between populations as well as strong sexual dimorphism in both body shape and armour were found. The lack of predator‐induced plasticity in development of the defence traits (viz. body armour and body depth) suggests that morphological anti‐predator traits in nine‐spined stickleback are strictly constitutive, rather than inducible. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Is phenotypic plasticity an explanation for the invasiveness of goldenrods (Solidago and Euthamia) in Europe?

High phenotypic plasticity contributes to invasiveness of alien species. Goldenrods of American origin (Solidago canadensis, S. gigantea and Euthamia graminifolia) have successfully invaded Europe, and this success can be related to their high phenotypic plasticity. The aim of our study was to test the hypothesis of higher phenotypic plasticity of invasive goldenrods by comparing them with native taxa: closely related Solidago virgaurea and similar ecologically, invasive in other regions Tanacetum vulgare. The species studied were grown in a common garden on three different substrates with different fertilizers. After 3 years, the height and number of ramets, biomass production and allocation, phenology and nitrate reductase activity (NRA) were measured. The highest level of phenotypic plasticity was exhibited by the competitively weak native species S. virgaurea. The invasive species produced relatively high biomass and exhibited a moderate level of phenotypic plasticity. Variability in the studied traits did not always correlate with differences in substrates. The invasion success of non-native goldenrods is not caused by their having a higher phenotypic plasticity than native species. Rather, the non-natives should be considered as “Jack of all traits, and master of some”.     

Life-history traits of non-native black bullhead Ameiurus melas with comments on its invasive potential

Life-history traits of non-native black bullhead Ameiurus melas from an oxbow of the River Váh (near the city of Komárno), located in the middle Danube reaches (Slovakia) were investigated during 2006 and 2007, including samples from the pre-spawning (March–May), spawning (May–June), post-spawning (late June–August) and resting (September) periods. In total, 600 specimens were examined. The sex ratio oscillated close to 1 : 1. Females were found to mature at 98.6–120 mm SL (118–145.1 mm TL, compared to 171–254 mm TL in the native North American populations). Spawning took place in May and June, although the spawning period in 2007 was more protracted. Based on the analyses of annual gonadal cycle and oocyte size distribution, the black bullhead is considered a multiple spawner with asynchronous oocyte development and indeterminate fecundity. Absolute fecundity ranged from 1238 to 4755 (mean 2610), and relative fecundity from 36.2 to 154.3 (mean 56.1) oocytes g⁻¹. The high flexibility in the life history of the black bullhead as demonstrated by its non-native populations, as well as its extreme tolerance and capability to live in systems with poor water quality, suggests that this species has a high potential to invade new areas and establish viable populations.     

Morphological divergence and flow‐induced phenotypic plasticity in a native fish from anthropogenically altered stream habitats

Understanding population‐level responses to human‐induced changes to habitats can elucidate the evolutionary consequences of rapid habitat alteration. Reservoirs constructed on streams expose stream fishes to novel selective pressures in these habitats. Assessing the drivers of trait divergence facilitated by these habitats will help identify evolutionary and ecological consequences of reservoir habitats. We tested for morphological divergence in a stream fish that occupies both stream and reservoir habitats. To assess contributions of genetic‐level differences and phenotypic plasticity induced by flow variation, we spawned and reared individuals from both habitats types in flow and no flow conditions. Body shape significantly and consistently diverged in reservoir habitats compared with streams; individuals from reservoirs were shallower bodied with smaller heads compared with individuals from streams. Significant population‐level differences in morphology persisted in offspring but morphological variation compared with field‐collected individuals was limited to the head region. Populations demonstrated dissimilar flow‐induced phenotypic plasticity when reared under flow, but phenotypic plasticity in response to flow variation was an unlikely explanation for observed phenotypic divergence in the field. Our results, together with previous investigations, suggest the environmental conditions currently thought to drive morphological change in reservoirs (i.e., predation and flow regimes) may not be the sole drivers of phenotypic change.     

Phenotypic plasticity of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> and <Emphasis Type="Italic">Phragmites australis</Emphasis> in response to nitrogen addition and intraspecific competition

Phenotypic plasticity of the two salt marsh grasses Spartina alterniflora and Phragmites australis in salt marshes is crucial to their invasive ability, but the importance of phenotypic plasticity, nitrogen levels, and intraspecific competition to the success of the two species is unclear at present. Spartina alterniflora Loisel. is an extensively invasive species that has increased dramatically in distribution and abundance on the Chinese and European coasts, and has had considerable ecological impacts in the regions where it has established. Meanwhile, Phragmites australis Cav., a native salt marsh species on the east coast of China, has replaced the native S. alterniflora in many marshes along the Atlantic Coast of the US. This study determined the effects of nitrogen availability and culm density on the morphology, growth, and biomass allocation traits of Spartina alterniflora and Phragmites australis. A large number of morphological, growth, and biomass parameters were measured, and various derived values (culm: root ratio, specific leaf area, etc.) were calculated, along with an index of phenotypic plasticity. Nitrogen addition significantly affected growth performance and biomass allocation traits of Spartina alterniflora, and culm density significantly affected morphological characteristics in a negative way, especially for Spartina alterniflora. However, there were no significant interactions between nitrogen levels and culm density on the morphological parameters, growth performances parameters, and biomass allocation parameters of the two species. Spartina alterniflora appears to respond more strongly to nitrogen than to culm density and this pattern of phenotypic plasticity appears to offer an expedition for successful invasion and displacement of Phramites australias in China. The implication of this study is that, in response to the environmental changes that are increasing nitrogen levels, the range of Spartina alterniflora is expected to continue to expand on the east coast of China.     

Shape plasticity in response to water velocity in the freshwater blenny Salaria fluviatilis

A non‐random association between an environmental factor and a given trait could be explained by directional selection (genetic determinism) and by phenotypic plasticity (environmental determinism). A previous study showed a significant relationship between morphology and water velocity in Salaria fluviatilis that conformed to functional expectations. The objective of this study was to test whether this relationship could be explained by phenotypic plasticity. Salaria fluviatilis from a Corsican stream were placed in four experimental channels with different water velocities (0, 10, 20 and 30 cm s^?1) to test whether there was a morphological response associated with this environmental factor. After 28 days, fish shape changed in response to water velocity without any significant growth. Fish in higher water velocities exhibited a more slender body shape and longer anal and caudal fins. These results indicate a high degree of morphological plasticity in riverine populations of S. fluviatilis and suggest that the previous relationship between morphology and water velocity observed in the field may largely be due to an environmental determinism.     

The effect of nitrogen and temperature changes on Solidago canadensis phenotypic plasticity and fitness

Phenotypic plasticity is commonly considered to contribute to the invasion success of invasive species. However, the importance of phenotypic plasticity, nitrogen (N) levels and warming to the invasion of invasive species is unclear. The effects of warming and N addition on the morphology, biomass allocation and biochemistry traits of Solidago canadensis and their plasticity were investigated by conducting a pot experiment. The results showed that the effect of N addition on biomass was improved for S. canadensis; whereas warming displayed no significant effect, their positive synergistic interact effect resulted in the overall significant increase in plant performance. The mean phenotypic plasticity indices (MPPI) of biochemistry and total parameters demonstrated a difference between operations, and the higher value was observed in N interaction with temperature treatments than N addition or warming alone. The observed MPPI indicated the biochemistry parameters > morphological parameters > allocation parameters. The MPPI of biochemistry parameters, morphological parameters and total parameters exhibited significant and positive correlations with N level and MPPI of morphological parameters was also significantly positively correlated with the fitness of S. canadensis. These results indicated that the global warming and N addition would make the invaded habitat more suitable for the growth of invasive S. canadensis, and even may effectively increase the invasion risk of S. canadensis through the enhanced phenotypic plasticity, which is a crucial factor to help species deal with the changing environment.     

Parallels,nonparallels, and plasticity in population differentiation of threespine stickleback within a lake

The frequent occurrence of parallel phenotypic divergence in similar habitats is often evoked when emphasizing the role of ecology in adaptive radiation and speciation. However, because phenotypic plasticity can contribute to the observed pattern of divergence, confirmation of divergence at loci underlying phenotypic traits is important for confirming adaptive divergence. In the present study, we examine parallel morphological, neutral, and potentially adaptive genetic divergence of threespine stickleback inhabiting different habitats within a lake. Three genetic clusters best explained the neutral genetic structure within the lake; however, morphological differences were only weakly connected to genetic clusters and there was considerable phenotypic variation within clusters. Among the factors that could contribute to the observed pattern of morphological and genetic divergence are phenotypic plasticity, selective mortality of hybrids, and habitat choice based on morphology. Several loci are identified as outliers indicating divergent selection between the morphs and some parallels in morphological and adaptive genetic divergence are found in stickleback spawning at two lava sites. However, neutral genetic structure indicates considerable genetic connectivity among the two lava sites, and the parallels in morphology may therefore represent selective distribution of phenotypes rather than parallel divergence. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 803–813.     

Leaf morphological plasticity in three dominant tree species in the Sundarbans mangrove forest of Bangladesh in different salinity zones

Mangrove tree species show plasticity in their leaf morphological traits in different salinity zones. However, leaf morphological plasticity and its causes in different salinity zones are incompletely understood. To understand the mechanism of plasticity, this study investigated the responses of three dominant tree species Sundri (Heritiera fomes), Gewa (Excoecaria agallocha) and Goran (Ceriops decandra) of the Sundarbans to the salinity gradients. A total of 17 leaf parameters were measured and quantified. All collected data were analyzed using univariate and multivariate statistical tools to investigate leaf morphological plasticity. A wide range of phenotypic plasticity was observed in all leaf parameters studied among the salinity zones of the Sundarbans. One-way ANOVA and Tukey’s posthoc test revealed significant differences (P?<?0.05) in all leaf parameters among the salinity zones and confirming that there was a high degree of phenotypic plasticity among the salinity zones of the Sundarbans. Petiole length (PL), leaf area (LA) and leaf length/petiole length (LL/PL) showed high level of plasticity among the salinity zones of the Sundarbans for each species of Sundri, Gewa and Goran. Plasticity index (PI) was developed in this study for each species studied. High level of phenotypic plasticity in these leaf traits reflects fitness of these species to different saline environments. Our results provide clear evidence that all the leaf parameters measured for three tree species viz., Sundri, Gewa and Goran effectively utilizes a plastic strategy in different salinity zones in the Sundarbans. Morphological trait plasticity could serve as powerful biological indicators to predict the shift of leaf morphology in upcoming environmental change events like sea level rise and reduction of fresh water flow from upstream.

     

Which Extent is Plasticity to Light Involved in the Ecotypic Differentiation of a Tree Species from Savanna and Forest?

Light intensity and heterogeneity are some of the main environmental factors that differ between forest and savanna habitats, and plant species from these habitats form distinct functional types. In this study, we tested the hypothesis that not only differences in morphological and physiological traits but also phenotypic plasticity in response to light are involved in adaptation to forest and savanna habitats by investigating ecotypic differentiation between populations of Plathymenia reticulata (Leguminosae: Mimosoideae), a tree from the Brazilian Atlantic Forest and the Brazilian Cerrado (savanna). Seeds from four natural populations (one from each biome core area and two from ecotonal regions) were grown in a common garden with four light treatments. Fifteen morphological and physiological characteristics were evaluated until individuals reached 6 mo old. Comparisons among populations showed differences for seven traits in at least one light treatment. These differences pointed to local adaptation to different biomes. Populations showed different levels of phenotypic plasticity in response to light in seven traits. Higher plasticity was found either in the forest core population or ecotonal populations; lower values were found in the cerrado core population. Lower plasticity in the cerrado population emphasizes the stress resistant syndrome, as lower plasticity is probably advantageous in a habitat where a conservative resource use is crucial. Higher plasticity in forest individuals suggests higher ability in exploiting the light heterogeneity in this habitat. Also, higher plasticity in ecotonal populations can be important to ensure the maintenance of P. reticulata in these temporally and spatially dynamic areas. Abstract in Portugese is available at http://www.blackwell‐synergy.com/loi/btp .