New and Old World phylogeography of pumpkinseed Lepomis gibbosus (Linnaeus, 1758): the North American origin of introduced populations in Europe

Hydrobiologia - To determine the origin of introduced European populations of pumpkinseed Lepomis gibbosus (Linnaeus, 1758)—a freshwater sunfish, native to eastern North America that has...     

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.     

Gradients of life-history variation: an intercontinental comparison of fishes

Multivariate analysis identified atwo-dimensional continuum of life-historyvariation among 301 fish species from Europe,North America, South America and the Atlanticand Pacific coasts of North America. The firstaxis was associated with larger body size,higher fecundity, delayed maturation, fewerreproductive events, and shorter breedingseason on one end and small size, lowfecundity, early maturity, multiplereproductive events per year, and prolongedbreeding season on the other. The second axiscontrasted fishes having larger eggs and moreparental care against fishes with the oppositesuite of traits.Phylogenetic affiliations of species wereapparent in the general patterns of ordinationof species within orders, indicatingevolutionary divergences in life-historypatterns. In fact, partitioning the variance oflife-history traits showed that taxonomic orderand latitude were the most important factorsand geographic region and habitat the least.Despite phylogenetic constraints, basiclife-history patterns showed consistencybetween distantly geographical regions,latitudinal ranges and basic adult habitats,indicating convergences in life-historypatterns. Although the basic life-historypatterns seemed repeatable among distantlyrelated taxa, geographical and latitudinalaffiliations were apparent. Species from SouthAmerica are skewed toward the opportunisticendpoint, whereas North American marine speciesare skewed toward the periodic endpoint of thetrilateral continuum model. Most of the fishspecies from South American data set came fromfluctuating environments, so an opportunisticstrategy of early maturation and continuousspawning permits efficient recolonization ofhabitats over small spatial scales. Incontrast, most species in the North Americanand European data sets came from seasonalhabitats that are nonetheless more hydrologicalstable, so a periodic strategy of delayingmaturation to attain large clutches enhancesadult survivorship during suboptimalenvironmental conditions and recruitment whenearly life stages encounter suitableenvironmental conditions. Similarly,latitudinal affiliations were also observed:opportunistic strategists more common intropical latitudes and periodic strategistsmore common in temperate and Arctic latitudes.     

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.     

Assessing fish metrics and biotic indices in a Mediterranean stream: effects of uncertain native status of fish

Implementation of the Water Framework Directive requires tools for measuring and monitoring the ecological status of aquatic ecosystems. Several indices are in use in the Iberian Peninsula, although there has been little comparison among them. We sampled the fish assemblage and limnological features of the Tordera stream (NE Spain) quarterly from September 2001 to May 2003 to evaluate the usefulness of several fish metrics and to compare habitat quality and biotic indices currently in use. Data for eight biotic and abiotic indices for this and three other Catalan river basins were also compiled in order to analyse the relationships among indices. In the Tordera stream, fish abundance and richness increased with stream order except in the last sampling site that had the lowest fish abundance owing to the effects of drought and water abstraction. Although most indices were positively correlated, some displayed low or null correlations particularly for the Tordera basin which is more affected by water abstraction and less by pollution; a commonly used physico-chemical index (ISQA) was the least correlated. In a regional fish index (IBICAT) under development, the brown trout (Salmo trutta) has been previously considered as introduced in the Tordera basin. Here, we report an old published record that demonstrates that trout was present before 1845 and we argue that its status should be considered as uncertain given the current information available. Whether brown trout is treated as native or introduced to this river basin has profound effects on the results of fish metrics because of its dominance in the upper reaches. We briefly discuss the role of introduced species, particularly in headwater streams, in the development of fish indices. Our study exemplifies the need for careful, basin-specific assessment of native/introduced status in the development of fish metrics. Handling editor: K. Martens     

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 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.     

Size-related effects and the influence of metabolic traits and morphology on swimming performance in fish

Energy metabolism fuels swimming and other biological processes. We compared the swimming performance and energy metabolism within and across eight freshwater fish species. Using swim tunnel respirometers, we measured the standard metabolic rate (SMR) and maximum metabolic rate (MMR) and calculated the critical swimming speed (U_crit). We accounted for body size, metabolic traits, and some morphometric ratios in an effort to understand the extent and underlying causes of variation. Body mass was largely the best predictor of swimming capacity and metabolic traits within species. Moreover, we found that predictive models using total length or SMR, in addition to body mass, significantly increased the explained variation of U_crit and MMR in certain fish species. These predictive models also underlined that, once body mass has been accounted for, U_crit can be independently affected by total length or MMR. This study exemplifies the utility of multiple regression models to assess within-species variability. At interspecific level, our results showed that variation in U_crit can partly be explained by the variation in the interrelated traits of MMR, fineness, and muscle ratios. Among the species studied, bleak Alburnus alburnus performed best in terms of swimming performance and efficiency. By contrast, pumpkinseed Lepomis gibbosus showed very poor swimming performance, but attained lower mass-specific cost of transport (MCOT) than some rheophilic species, possibly reflecting a cost reduction strategy to compensate for hydrodynamic disadvantages. In conclusion, this study provides insight into the key factors influencing the swimming performance of fish at both intra- and interspecific levels.     

Morphological variation in pumpkinseed Lepomis gibbosus introduced into Iberian lakes and reservoirs; adaptations to habitat type and diet?

The morphology of pumpkinseed Lepomis gibbosus populations from five Catalonian waterbodies (north-eastern Spain) that vary in hydromorphometry was examined and compared to a native North American reference site that contained two morphological variants of this species. Populations exhibited significant differences in fin location, body depth and caudal peduncle length, which are known to have functional significance in the hydrodynamics of swimming and hence the foraging mode. Differences were also noted in internal morphological traits functionally related to prey selection. Pumpkinseed populations that fed extensively on zooplankton showed narrow gill raker spacing, and mollusc-feeding populations had longer and wider pharyngeal bones. Discriminant function analysis (DFA) provided significant separation of all populations on the basis of both external and internal morphology, with the main axis of separation being geographical rather than environmental. The secondary DFA axis, however, did separate populations that fed primarily on zooplankton from those that were primarily benthic invertebrate feeders. In this regard, a population that occupied an Iberian steep-sided reservoir with an unstable littoral zone showed similar morphological adaptations to the limnetic morphological variant native in North America, supporting previous studies showing that fish morphology is strongly affected by prey type and feeding mode. The results suggest that pumpkinseeds are able to adapt, morphologically, to the types of habitats and prey present in Catalonian waterbodies, and this may partially explain why they are so successful in areas where they have been introduced.     

To be,or not to be,a non‐native freshwater fish?

We examine the evolving concept of what constitutes a non‐native (or alien) freshwater fish. In an attempt to distinguish between biogeographical and socio‐political perspectives, we review the patterns in the introduction and dispersal of non‐native fishes in Europe and North America, and especially the recent expansion of Ponto‐Caspian gobies in Europe. We assess patterns in the development of national policy and legislation in response to the perceived threat of non‐native fish introductions to native species and ecosystems. We review, and provide a glossary of, the terms and definitions associated with non‐native species. Finally, we discuss perspectives as regards the future treatment of naturalized species.