Patterns of morphological variation among native and non-native pumpkinseed (<Emphasis Type="Italic">Lepomis gibbosus</Emphasis>) populations: shared and unique aspects of diversification

Contemporary patterns of morphological variation among populations reflects the interplay between historic and contemporary processes that result from selection and constraint. Using the pumpkinseed (Lepomis gibbosus), a species native to North America and introduced to Europe, we assessed the shared and unique aspects of morphological divergence in lentic and lotic environments among native and non-native populations. Ten native and thirteen non-native pumpkinseed populations were collected between 2003 and 2010 from lakes, rivers and reservoirs within the Iberian Peninsula and east-central North America. Fifteen linear external measurements among homologous landmarks that pertain to body size, fin position and fin size were taken from all sampled individuals. Eleven of these measurements were used to test for morphological differences among populations. Pumpkinseed found in lotic water bodies exhibited a more anterior placement of pectoral and pelvic fins and a deeper caudal peduncle and body than those found in lentic water bodies from the same geographic region. However, pumpkinseed also showed morphological differences between geographic origins: pumpkinseed from native populations exhibit a more posterior placement of pectoral and pelvic fins, a narrower anterior caudal peduncle and a more slender body than pumpkinseed from non-native populations. In addition, unique responses of populations to waterbodies within geographic origins revealed a shift between water body types that was opposite in direction for native and non-native populations. Native populations exhibited shorter and deeper caudal peduncles and deeper bodies in lotic habitats, whereas non-native populations showed longer and slender caudal peduncles and more slender bodies in the same type of habitat. Our study demonstrates that contemporary patterns of morphological variation among native and non-native pumpkinseed populations can be explained by contemporary selection and/or a common plastic developmental response among water bodies, historical effects related to geographic origin and unique responses of populations to habitats within geographic origin, and that the effects of history and the interaction between history and contemporary habitat were larger than contemporary processes in explaining morphological variation at this large spatial scale.     

Life-history traits of introduced Iberian pumpkinseed Lepomis gibbosus relative to native populations. Can differences explain colonization success?

To assess the colonization success of pumpkinseed Lepomis gibbosus on the Iberian Peninsula, life-history attributes of pumpkinseed populations from Spanish water bodies were compared to populations in the northern and southern parts of its native range, as well as to those of English water bodies where introduced populations have shown minimal natural range expansion. Discriminate function analysis using five population characteristics [mean age at maturity, mean total length ( L _T) at maturity, gonado-somatic index ( I _G), mean L _T at age 2 years as an indication of juvenile growth rate and relative body condition] strongly differentiated populations from the four regional study areas. Spanish populations were early maturing, showed moderate juvenile growth rate and L _T at maturity and high I _G relative to the other population groups. Spanish populations matured significantly earlier than Canadian and English populations, but not southern U.S.A. populations. Spanish populations, however, had a significantly higher I _G than southern U.S.A. populations, suggesting greater reproductive output. Considering these differences in the context of the Winemiller and Rose triangular life-history strategy model, the pumpkinseed exhibits 'equilibrium' life-history traits that have made other non-native fishes successful invaders in areas where habitat alteration has resulted in more predictable discharge regimes and water levels. Populations in Iberia, however, appear to have achieved their success by adopting a more 'opportunistic' life-history strategy than their native counterparts. High levels of life-history plasticity appear to contribute to the success of this species on the Iberian Peninsula.     

Old world versus new world: life-history alterations in a successful invader introduced across Europe

We examined differences in pumpkinseed (Lepomis gibbosus) life-history traits between native North American and introduced European populations, and tested three life-history predictions related to the effect of temperature, growth, waterbody size, and the presence/absence of predators on native and non-native populations. Pumpkinseed populations exhibit more ‘opportunistic’ traits (earlier maturity, smaller size at maturity, and higher reproductive allocation) in their introduced European range than those in their native range. Predictions of life-history traits were improved when indicators of juvenile growth rate (mean length at age 2), waterbody size (surface area), and thermal regime (air temperature degree-days above 10 °C) were incorporated into models along with continental location, but European pumpkinseed populations exhibit more opportunistic life-history traits than North American populations even when these factors are accounted for. Native pumpkinseed in waterbodies containing piscivores mature later and at a larger size, and have lower gonadosomatic indices than those in waterbodies lacking piscivores, whereas there is no significant difference in the same three life-history traits between European waterbodies containing or lacking piscivores. Because congeneric competitors of the pumpkinseed are absent from Europe, the apparent absence of a predator life-history effect there could also be due to the absence of the major sunfish competitors. In either case, the evolution and maintenance of more opportunistic traits in European pumpkinseed can likely be attributed to enemy release, and this may explain the successful establishment and spread of pumpkinseed in many parts of Europe.     

Morphological change and phenotypic plasticity in native and non‐native pumpkinseed sunfish in response to sustained water velocities

Phenotypic plasticity can contribute to the proliferation and invasion success of nonindigenous species by promoting phenotypic changes that increase fitness, facilitate range expansion and improve survival. In this study, differences in phenotypic plasticity were investigated using young‐of‐year pumpkinseed sunfish from colonies established with lentic and lotic populations originating in Canada (native) and Spain (non‐native). Individuals were subjected to static and flowing water treatments for 80 days. Inter‐ and intra‐population differences were tested using ancova and discriminant function analysis, and differences in phenotypic plasticity were tested through a manova of discriminant function scores. Differences between Iberian and North American populations were observed in dorsal fin length, pectoral fin position and caudal peduncle length. Phenotypic plasticity had less influence on morphology than genetic factors, regardless of population origin. Contrary to predictions, Iberian pumpkinseed exhibited lower levels of phenotypic plasticity than native populations, suggesting that canalization may have occurred in the non‐native populations during the processes of introduction and range expansion.     

Profiling invasive fish species: the importance of phylogeny and human use

Understanding the ecological differences between native and invasive species is of considerable scientific and practical interest. We examined such differences between native and invasive inland fish species from the Iberian Peninsula in order to analyse the importance of phylogenetic correction and variability (in addition to central tendency). We collected 26 quantitative and qualitative variables on the ecology, life‐history traits and human use of the 69 inland fish species of the Iberian Peninsula, including native, invasive and migratory species. The taxonomic distribution of invasive fish species deviated significantly from world freshwater richness and in contrast to native species, invasive fish belongs to only five taxonomic orders but to a wide spectrum of families not native to the Iberian Peninsula. Because the life‐history traits were highly dependent on taxonomy, the results, with or without applying phylogenetic methods, differed and after accounting for phylogeny, invasive species displayed higher and wider latitude in general and a different reproductive season mainly among salmonids and cyprinids. Human use was also significantly different between native and invasive fish species and produced more variability in life‐history traits of invasive species and uneven taxonomic distribution because of the high diversity of species introduced. We show that accounting for taxonomy and studying variability in addition to central tendency is important in the comparison of life‐history traits between native and invasive species.     

Origin matters: widely distributed native and non-native species benefit from different functional traits

Recently, ecologists debated whether distinguishing native from non-native species is sensible or not. One argument is that widespread and less widespread species are functionally different, whether or not they are native. An opposing statement points out ecologically relevant differences between native and non-native species. We studied the functional traits that drive native and non-native vascular plant species frequency in Germany by explaining species grid-cell frequency using traits and their interaction with status. Native and non-native species frequency was equally driven by life span, ploidy type and self-compatibility. Non-native species frequency rose with later flowering cessation date, whereas this relationship was absent for native species. Native and non-native species differed in storage organs and in the number of environmental conditions they tolerate. We infer that environmental filters drive trait convergence of native and non-native species, whereas competition drives trait divergence. Meanwhile, introduction pathways functionally bias the frequency of non-native species.     

Geographic variation in floral morphology and style-morph ratios in a sexually polymorphic daffodil

Narcissus triandrus is a tristylous daffodil from the Iberian Peninsula that has striking geographical variation in floral morphology and style-morph ratios. Here, we investigate the relation between this variation and ecological factors to understand mechanisms governing morph ratios. We estimated morph ratios in 124 populations throughout the range of N. triandrus and measured 13 morphological traits in 35-78 populations. Sampling of morph ratios conducted over 2-10-yr intervals demonstrated strong temporal stability. Variation in floral and vegetative traits enabled statistical prediction of morph-frequency variation among populations. Latitudinal gradients in precipitation and temperature were correlated with plant and flower size, with larger flowers in northern populations associated with bumblebee visitation and stylar dimorphism. Flowers of the L- and M-morphs differed significantly from the S-morph in several size-related characters, unlike other tristylous species. This pattern and the similarity in anther positions of the L- and M-morphs suggest that the M-morph of N. triandrus originated through genetic modifiers that shortened styles of the L-morph. Our findings support the hypothesis that geographical variation in style-morph ratios is largely governed by climatic gradients in the Iberian Peninsula, which influence the floral morphology and pollination biology of N. triandrus populations.     

An Anthropogenic Habitat Facilitates the Establishment of Non-Native Birds by Providing Underexploited Resources

Anthropogenic modification of habitats may reduce the resources available for native species, leading to population declines and extinction. These same habitats often have the highest richness of non-native species. This pattern may be explained if recently human-modified habitats provide novel resources that are more accessible to non-native species than native species. Using non-native birds in the Iberian Peninsula as a case study, we conduct a large-scale study to investigate whether non-native species are positively associated with human modified habitats, and to investigate whether this positive association may be driven by the presence of resources that are not fully exploited by native species. We do this by comparing the functional diversity and resource use of native and non-native bird communities in a recently human-modified habitat (rice fields) and in more traditional habitats in the Iberian Peninsula. The functional diversity of native bird communities was lower in rice fields, but non-native birds were positively associated with rice fields and plugged this gap. Differences in resource use between native and non-native species allowed non-native species to exploit resources that were plentiful in rice fields, supporting the role of underexploited resources in driving the positive association of non-native birds with rice fields. Our results provide a potential mechanism explaining the positive association of non-native species with anthropogenic habitats, and further work is needed to test if this applies more generally.     

Life-history traits of invasive fish in small Mediterranean streams

We compared the life-history traits of native and invasive fish species from Catalan streams in order to identify the characters of successful invasive fish species. Most of the exotic fish species were characterized by large size, long longevity, late maturity, high fecundity, few spawnings per year, and short reproductive span, whereas Iberian native species exhibited predominantly the opposite suite of traits. Species native to the southeastern Pyrenees watershed were also significantly different from species native to the rest of the Iberian Peninsula but not native to this watershed. Iberian exotic species come predominantly from large river basins, whereas Catalan streams (and other small, coastal river basins) correspond to basins and streams of a smaller size and different hydrology, with differences in species composition and life-history traits of fish. The occurrence and spread of invasive species was not significantly related to life-history traits but to introduction date. The successful prediction of future invasive species is limited due to small differences in life-history and ecological traits between native and exotic species. Fecundity, age at maturity, water quality flexibility, tolerance to pollution and habitat seem the most discriminating life-history variables. This revised version was published online in July 2006 with corrections to the Cover Date.     

Distance between south-European and south-west Asiatic refugial areas involved morphological differentiation: Pinus sylvestris case study

The phenotypic differentiation of relic P. sylvestris in southern Europe and southwestern Asia was verified using thirty-two populations sampled from the Iberian Peninsula, Massif Central, Balkan Peninsula, Crimea and Anatolia. Twenty-one morphological and anatomical needle traits and 18 cone morphological characteristics were examined to describe the population diversity and differentiation. The needle characters were not correlated to those of cone. The differences between regions were significant based on 12 needle and 9 cone characteristics, suggesting spatial isolation. The differentiation between the Iberian and Anatolian populations was the highest, which indicates the isolation by distance. The high level of morphological differentiation was also found among Iberian populations, supporting the already known complex history of the species in that region. Populations within other regions were differentiated at lower levels; however, the West Anatolian populations differed morphologically from the eastern ones. The described pattern of morphological differentiation supports the idea of the long-lasting existence of P. sylvestris in the south-European and Anatolian mountain regions. To conserve this variation, seed transfer between regions in the forest economy should be restricted.     

Invasive and Non-Invasive Congeners Show Similar Trait Shifts between Their Same Native and Non-Native Ranges

Differences in morphological or ecological traits expressed by exotic species between their native and non-native ranges are often interpreted as evidence for adaptation to new conditions in the non-native ranges. In turn this adaptation is often hypothesized to contribute to the successful invasion of these species. There is good evidence for rapid evolution by many exotic invasives, but the extent to which these evolutionary changes actually drive invasiveness is unclear. One approach to resolving the relationship between adaptive responses and successful invasion is to compare traits between populations from the native and non-native ranges for both exotic invaders and congeners that are exotic but not invasive. We compared a suite of morphological traits that are commonly tested in the literature in the context of invasion for three very closely related species of Centaurea, all of which are sympatric in the same native and non-native ranges in Europe and North America. Of these, C. solstitialis is highly invasive whereas C. calcitrapa and C. sulphurea are not. For all three species, plants from non-native populations showed similar shifts in key traits that have been identified in other studies as important putative adaptive responses to post-introduction invasion. For example, for all three species plants from populations in non-native ranges were (i) larger and (ii) produced seeds that germinated at higher rates. In fact, the non-invasive C. calcitrapa showed the strongest trait shift between ranges. Centaurea solstitialis was the only species for which plants from the non-native range increased allocation to defensive spines, and allocated proportionally less resources to reproduction, patterns contrary to what would be predicted by theory and other empirical studies to enhance invasion. Our results suggest caution when interpreting the commonly observed increase in size and reproductive capacity as factors that cause exotics to become invaders.     

Trophic polymorphism in introduced pumpkinseed (<Emphasis Type="Italic">Lepomis gibbosus</Emphasis>) inhabiting Iberian reservoirs

Introduced pumpkinseed in Iberian reservoirs display marked external morphological differentiation along two simultaneous dimensions of flow and trophic structure. We assessed the degree of internal morphological differentiation using gill rakers, pharyngeal jaws and the levator posterior muscle among pumpkinseeds occupying four different habitats and determined whether prey consumption accounted for any discernible differences in feeding structures among ecomorphs. Results showed significant differentiation by habitat based on pharyngeal muscle and jaw dimensions in all study reservoirs, with pelagic pumpkinseeds having smaller jaws than littoral pumpkinseeds in four of the five reservoirs. Gill rakers, however, differentiated morphs in only three of the five reservoirs, corresponding to differences in zooplankton consumption among pelagic and littoral individuals in those reservoirs. Based on all internal morphological traits, greater divergence was seen along the littoral-pelagic trophic axis in the lacustrine zones of reservoirs compared to the fluvial zone. Overall differences noted in internal morphology are likely the result of phenotypic plasticity; the ability of this species to readily adapt to changing physical environments may explain the success of the pumpkinseed in its introduced range.     

Root architecture,plant size and soil nutrient variation in natural populations of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Phenotypic variation in ecologically important traits may vary at large and small geographic scales, and may be shaped by natural selection. Here our explicit aim is to evaluate phenotypic differentiation among local populations and examine its relationship with ecological edaphic and climatic features that could lead to local adaptation. We characterized six populations of the model plant Arabidopsis thaliana over 3 years in the field in its native range. At each site, we measured edaphic conditions and aboveground and belowground phenotypes. In addition, we grew plants from the six characterized populations in a common greenhouse along with an additional fifteen populations from the Iberian Peninsula to examine evolutionary and ecological differentiation among populations, and relationships between geographic and ecological distance to phenotypic differences among populations. Significant differences in aboveground and belowground traits, population density, and micro- and macro-nutrient soil concentrations were found among the field populations. In particular, root architectural traits differed significantly among field populations. Complex patterns of ecological differences among population and plant phenotypes emerged when examining edaphic conditions in the Extremadura region, and geographic and climate variables at a broader scale of the Iberian Peninsula. We report levels of phenotypic variation at the local scale comparable to those found at broad geographic scales and report that local edaphic conditions contribute to population-level phenotypic variation in root and shoot traits. To our knowledge, these are the first reports of among population root architectural variation from natural field populations for this model organism. We demonstrate how ecological features, such as soil nutrients, can be associated with the phenotypic variation of A. thaliana measured in natural populations and may contribute to adaptive differentiation at a local scale.     

Morphological variation between non‐native lake‐ and stream‐dwelling pumpkinseed Lepomis gibbosusin the Iberian Peninsula

The objective of this study was to test if morphological differences in pumpkinseed Lepomis gibbosus found in their native range (eastern North America) that are linked to feeding regime, competition with other species, hydrodynamic forces and habitat were also found among stream‐ and lake‐ or reservoir‐dwelling fish in Iberian systems. The species has been introduced into these systems, expanding its range, and is presumably well adapted to freshwater Iberian Peninsula ecosystems. The results show a consistent pattern for size of lateral fins, with L. gibbosus that inhabit streams in the Iberian Peninsula having longer lateral fins than those inhabiting reservoirs or lakes. Differences in fin placement, body depth and caudal peduncle dimensions do not differentiate populations of L. gibbosus from lentic and lotic water bodies and, therefore, are not consistent with functional expectations. Lepomis gibbosus from lotic and lentic habitats also do not show a consistent pattern of internal morphological differentiation, probably due to the lack of lotic–lentic differences in prey type. Overall, the univariate and multivariate analyses show that most of the external and internal morphological characters that vary among populations do not differentiate lotic from lentic Iberian populations. The lack of expected differences may be a consequence of the high seasonal flow variation in Mediterranean streams, and the resultant low‐ or no‐flow conditions during periods of summer drought.     

Biogeographic differences between native and non-native populations of crayfish alter species coexistence and trophic interactions in mesocosms

Biogeographical comparisons of native and non-native populations allow researchers to understand the degree to which traits contributing to invasion success are intrinsic or change during the invasion process. Here, we investigate whether traits underlying interspecific competition change following invasion and whether these alter the impacts of two crayfish congeners that have invaded into each other’s native ranges. Specifically, we compared native and non-native populations of rusty (Faxonius rusticus) and virile crayfish (F. virilis). We compared native and non-native populations of each species using laboratory assays to examine aggression and large mesocosms with the congeners in sympatry to examine growth and survival as well as impacts on lower trophic levels. We found that non-native virile crayfish were more aggressive in response to a threat than native virile crayfish and exhibited greater growth and survival in sympatry with rusty crayfish. These intraspecific differences were large enough to alter coexistence between species in the mesocosm experiment, which is consistent with patterns of coexistence between these species in the field. We did not observe differences in traits between native and non-native rusty crayfish, but rusty crayfish were consistently competitively dominant over virile crayfish in paired laboratory assays. Non-native populations of both species had greater impacts on lower trophic levels than native populations. Taken together, these findings provide new evidence that trait changes during invasions may enhance ecological impacts of invasive animals and their ability to compete with closely related native species.     

Morphological differentiation in introduced pumpkinseed Lepomis gibbosus (L.) occupying different habitat zones in Portuguese reservoirs

Introduced pumpkinseed Lepomis gibbosus sampled from four habitat zones (fluvial pelagic, fluvial littoral, lacustrine pelagic and lacustrine littoral) in three Portuguese reservoirs were used to test the hypotheses that habitats with the least similar characteristics will show the most differentiation, and that morphological differences will relate to functional adaptations to flow and trophic habitats. Results from discriminant function analysis and ANCOVA showed that there were significant differences in external morphology in pumpkinseed captured from the four habitat zones in all three reservoirs. Littoral and pelagic differentiation was stronger than fluvial and lacustrine differentiation in all of the reservoirs, and the most significant variable that differentiated pumpkinseed from the littoral and pelagic habitats was body depth. The illustration of external morphological differentiation in pumpkinseed along both habitat dimensions demonstrates the high degree of morphological plasticity of this introduced species.     

Reproductive habitat selection in alien and native populations of the genus Discoglossus

The existence of suitable breeding habitats is an important factor explaining the regional presence of an anuran species. This study examined patterns of habitat selection in populations of three species of the genus Discoglossus: Discoglossus galganoi (south-western Iberian Peninsula), Discoglossus scovazzi (Morocco) and Discoglossus pictus (three different areas were included in the study: Sicily, Tunisia and north-eastern Iberian Peninsula). The populations of D. pictus on the Iberian Peninsula are allochthonous, and analysis of these patterns may provide insights into the processes that regulate the invasion phase. The hypotheses tested were: (i) congeneric species show the same patterns of habitat selection, and alien species have been established following these patterns; (ii) there are differences in species associations between assemblages structured deterministically and by chance, i.e. native versus invaded assemblages. The larval habitats of three species of this genus were characterized by measuring physical and chemical parameters of the water bodies. We examined the covariation between the presence of Discoglossus species and the species richness of sympatric anurans, and investigated a possible relationship between morphological similarity (as a proxy of functional group) and overlap in habitat use. The results showed that congeneric species are morphologically conservative and also select very similar types of aquatic habitat. The alien population and other sympatric species showed a high degree of overlap in habitat use, which was greater than that observed in the native assemblage with a similar functional richness. Species associations were not structured on the basis of morphological similarity in any of the assemblages. Among native populations, the presence of Discoglossus was either negatively correlated or not significantly correlated with species richness. Only the alien population showed a positive correlation between its presence and species richness, which suggests a loss of assemblage structure.     

Introduced plants of Lupinus polyphyllus are larger but flower less frequently than conspecifics from the native range: Results of the first year

Introduced species, which establish in novel environments, provide an opportunity to explore trait evolution and how it may contribute to the distribution and spread of species. Here, we explore trait changes of the perennial herb Lupinus polyphyllus based on 11 native populations in the western USA and 17 introduced populations in Finland. More specifically, we investigated whether introduced populations outperformed native populations in traits measured in situ (seed mass) and under common garden conditions during their first year (plant size, flowering probability, and number of flowering shoots). We also explored whether climate of origin (temperature) influenced plant traits and quantified the degree to which trait variability was explained collectively by country and temperature as compared to other population‐level differences. Three out of four plant traits differed between the native and introduced populations; only seed mass was similar between countries, with most of its variation attributed to other sources of intraspecific variation not accounted for by country and temperature. Under common garden conditions, plants originating from introduced populations were larger than those originating from native populations. However, plants from the introduced range flowered less frequently and had fewer flowering shoots than their native‐range counterparts. Temperature of a population''s origin influenced plant size in the common garden, with plant size increasing with increasing mean annual temperature in both native and introduced populations. Our results of the first year reveal genetic basis for phenotypic differences in some fitness‐related traits between the native and introduced populations of L. polyphyllus. However, not all of these trait differences necessarily contribute to the invasion success of the species and thus may not be adaptive, which raises a question how persistent the trait differences observed in the first year are later in individuals’ life for perennial herbs.     

Life‐history variability of non‐native centrarchids in regulated river systems of the lower River Guadiana drainage (south‐west Iberian Peninsula)

Life‐history variability of two non‐native centrarchids, pumpkinseed Lepomis gibbosus and largemouth bass Micropterus salmoides, was evaluated in three stream stretches of the lower River Guadiana drainage (south‐west Iberian Peninsula) with different degrees of regulated flows. Abundance, condition and population structure differed among populations for both species, but invasion success was lower in the least regulated river. Lepomis gibbosus were abundant and had multiple age classes in the three river sites, whereas M. salmoides were less abundant and mainly represented by young‐of‐the‐year fish. Juvenile growth in L. gibbosus was similar in all three populations, though longevity was slightly greater in the population from the River Guadiana mainstream. Lepomis gibbosus exhibited a long reproductive season, but the duration of season, size at maturity and reproductive effort varied among populations. The life‐history differences found demonstrate the importance of species adaptation to local conditions which might favour their invasion success. Lepomis gibbosus were more adaptable and resilient to local conditions, whereas M. salmoides seemed dependent on reservoirs and large rivers for maintenance of riverine populations.     

Effects of the non-native amphibian species Discoglossus pictus on the recipient amphibian community: niche overlap, competition and community organization

The painted frog, Discoglossus pictus, was introduced into the Iberian Peninsula 100 years ago and its distribution has steadily increased since then. We studied the effects of this non-native amphibian species on the native ones in the northeastern area of the peninsula. We compared amphibian assemblages in regions with and without D. pictus to estimate niche overlap between species. Additionally, we carried out a laboratory evaluation of the effects of competition between the non-native and the two native species with which it overlaps most commonly: Bufo calamita and Pelodytes punctatus. The presence of D. pictus larvae reduced the survival, body mass and activity of B. calamita, and increased time to metamorphosis. Furthermore, D. pictus showed the highest consumption rate while P. punctatus showed the lowest. One possible consequence of these competitive interactions is an alteration of species co-occurrence patterns in the recipient community on a regional scale. In the non-invaded area, the checkerboard score (C-score) of co-occurrence indicated that the community was structured, whereas the standardized C-score in the invaded area indicated that the community did not differ significantly from having a random structure. These results suggest that competition between native and non-native species can cause recipient communities to become less structured.