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.     

External morphology of a Slovenian population of pumpkinseed Lepomis gibbosus (L.) from a habitat with extreme thermal conditions

The external morphology and growth variability of morphometric characters of pumpkinseed (n = 141) from an oxbow (River Sava, Slovenia) that receives thermal effluent were examined using triple regression analysis. Differences in external morphology between pumpkinseed from the oxbow and both native North American (River Otonabee, Looncall Lake, Canada) and non‐native Central European populations (side arms of the River Danube, Slovakia) were evaluated. Two possible morphotypes among adults were observed, whereas the morphology of juveniles appears rather uniform across geographical location (i.e. Otonabee, Looncall, Danube). This suggests that environmental conditions, i.e. epigenetical information, seem to be responsible for most of variability in pumpkinseed morphology, which represents a function of epigenetical mechanisms. However, further investigation into epigenetical interactions, especially early development, fecundity, number of spawning acts per season, parental care, egg size and age at maturation are necessary to test this hypothesis.     

Monogenean parasites of introduced pumpkinseed Lepomis gibbosus (Centrarchidae) in the Danube River Basin

The pumpkinseed, Lepomis gibbosus, a centrarchid fish native to eastern North America, was introduced into Europe at the end of the 19th century. Lepomis gibbosus now represents one of the most successful of introduced fish species, having spread widely throughout many European countries. In this study, we collected monogenean parasites of L. gibbosus from four sites in the Danube River Basin. We recorded four monogenean species: Gyrodactylus sp., probably acquired in the area of introduction, and Onchocleidus similis, O. dispar and Actinocleidus recurvatus, species introduced together with their host. Onchocleidus similis dominated the parasite community at all sampling sites. Actinocleidus recurvatus has only been reported from southern Europe to date and, therefore, this is the first report of A. recurvatus from Central Europe and the Danube River Basin. We observed high morphological variability in both anchor pairs of A. recurvatus. Only in O. similis was there a positive correlation observed between haptor metric traits and fish host length. Differences in metric traits between native and introduced O. similis populations may reflect a difference in fish host size, the introduced fish host achieving a smaller size than fish from native populations. A detailed analysis of microhabitat distribution indicated a preference for the second gill arch for both dominant species O. similis and A. recurvatus, with dissimilar site segregation.     

Tree-based ensembles unveil the microhabitat suitability for the invasive bleak (Alburnus alburnus L.) and pumpkinseed (Lepomis gibbosus L.): Introducing XGBoost to eco-informatics

Random Forests (RFs) and Gradient Boosting Machines (GBMs) are popular approaches for habitat suitability modelling in environmental flow assessment. However, both present some limitations theoretically solved by alternative tree-based ensemble techniques (e.g. conditional RFs or oblique RFs). Among them, eXtreme Gradient Boosting machines (XGBoost) has proven to be another promising technique that mixes subroutines developed for RFs and GBMs. To inspect the capabilities of these alternative techniques, RFs and GBMs were compared with: conditional RFs, oblique RFs and XGBoost by modelling, at the micro-scale, the habitat suitability for the invasive bleak (Alburnus alburnus L.) and pumpkinseed (Lepomis gibbosus L.). XGBoost outperformed the other approaches, particularly conditional and oblique RFs, although there were no statistical differences with standard RFs and GBMs. The partial dependence plots highlighted the lacustrine origins of pumpkinseed and the preference for lentic habitats of bleak. However, the latter depicted a larger tolerance for rapid microhabitats found in run-type river segments, which is likely to hinder the management of flow regimes to control its invasion. The difference in the computational burden and, especially, the characteristics of datasets on microhabitat use (low data prevalence and high overlapping between categories) led us to conclude that, in the short term, XGBoost is not destined to replace properly optimised RFs and GBMs in the process of habitat suitability modelling at the micro-scale.     

Estimating diet in individual pumpkinseed sunfish Lepomis gibbosus using stomach contents,stable isotopes and parasites

The diets of 99 pumpkinseed sunfish Lepomis gibbosus from a pair of small, adjacent lakes in Ontario, Canada, were estimated from their stomach contents, trophically transmitted parasites and stable isotopes of carbon and nitrogen in fish tissue. The three methods provided virtually unrelated information. There was no significant correlation in the importance of any prey item across all three methods. Fish with similar diets according to one method of estimating diet showed no tendency to be similar according to other methods. Although there was limited variation in fish size and the spatial scale of the study was small, both fish size and spatial origin showed comparatively strong associations with diet data obtained with all three methods. These results suggest that a multidisciplinary approach that accounts for fish size and spatial origins is necessary to accurately characterize diets of individual fish.     

Induction of fever in a teleost fish (Lepomis gibbosus, L.)

Although fishes are ectotherms they are nevertheless able to thermoregulate behaviorally by selecting appropriate water temperatures (1). In a temperature gradient fish will congregate to a species-specific range of preferred temperature (“final thermal referendum”) which is unaffected by previous thermal history of the individual (2,3). Several aquatic (and terrestrial) ectothermic vertebrates have been found to exhibit “behavioral fever” which is manifested as an increase in preferred temperature above the final thermal preferendum (4). Fever can be elicited by pyrogens: whole bacteria (alive or killed), components of bacterial cellwall (endotoxins), endogenous pyrogens, prostaglandins or from several other sources (5). Since the results with fever induction in fish using whole bacteria or endotoxins are very scarce the aim of the present work was to compare possible thermoregulatory effects of endotoxins and prostaglandins in the same species (Lepomis gibbosus, L.) by means of identical methods.     

Foraging performance of diet-induced morphotypes in pumpkinseed sunfish (Lepomis gibbosus) favours resource polymorphism

Morphological plasticity can influence adaptive divergence when it affects fitness components such as foraging performance. We induced morphological variation in pumpkinseed sunfish (Lepomis gibbosus) ecomorphs and tested for effects on foraging performance. Young-of-year pumpkinseed sunfish from littoral and pelagic lake habitats were reared each on a 'specialist diet' representing their native habitat-specific prey, or a 'generalist diet' reflecting a combination of native and non-native prey. Specialist and generalist diets, respectively, induced divergent and intermediate body forms. Specialists had the highest capture success on their native prey whereas generalist forms were inferior. Specialists faced trade-offs across prey types. However, pelagic specialists also had the highest intake rate on both prey types suggesting that foraging trade-offs are relaxed when prey are abundant. This increases the likelihood of a resource polymorphism because the specialized pelagic form can be favoured by directional selection when prey are abundant and by diversifying selection when prey resources are restricted.     

Life-history traits and potential invasiveness of introduced pumpkinseed Lepomis gibbosus populations in northwestern Europe

To evaluate the potential invasiveness of pumpkinseed Lepomis gibbosus introduced to northwestern European inland waters, growth and reproduction traits were examined in ten populations along a trajectory spanning northwestern Europe (Norway, England, Holland, Belgium and France) and evaluated in light of published dataset from Europe. In the 848 pumpkinseed captured, maximum age was 3–4 years, with a sex ratio near unity in all but one population. Significant variations with increasing latitude were observed in adult growth (age 2–3 increment in total length, TL) and mean age at maturity (A _M), with non-significant variations observed in juvenile growth (TL at age 2), sex ratio and gonado-somatic index. As observed elsewhere in Europe, mean A _M decreased significantly with increasing TL at age 2. Using this relationship, which has been proposed elsewhere as a potential predictive model of pumpkinseed invasiveness, eight of the ten populations could be provisionally categorized as ‘non-invasive’ (five populations), ‘transitional’ (one population) and ‘potentially invasive’ (two populations), with two populations not categorized due to insufficient data. Based on the available knowledge on each population, the relationship between juvenile growth and age at maturity appeared to predict reasonably the status of pumpkinseed in northwestern Europe and its applicability to other species should be tested.     

Flexible mate choice may contribute to ecotype assortative mating in pumpkinseed sunfish (Lepomis gibbosus)

Gene flow is expected to limit adaptive divergence, but the ecological and behavioural factors that govern gene flow are still poorly understood, particularly at the earliest stages of population divergence. Reduced gene flow through mate choice (sexual isolation) can evolve even under conditions of subtle population divergence if intermediate phenotypes have reduced fitness. We indirectly tested the hypothesis that mate choice has evolved between coexisting littoral and pelagic ecotypes of polyphenic pumpkinseed sunfish (Lepomis gibbosus) that have diverged in morphology and resource use and where intermediate phenotypes have reduced performance. We assessed the ecotype of nesting males and females using stable isotope estimates of diet and a divergent male morphological trait, oral jaw width. We found positive assortative mating between ecotypes in a common spawning habitat along exposed lake shorelines, but contrary to expectations, assortative mating was variably expressed between two sampling years. Although the factors that influence variable assortative mating remain unclear, our results are consistent with mate choice being expressed by ecotypes. Despite being variably expressed, mate choice will reduce gene flow between ecotypes and could contribute to further adaptive divergence depending on its frequency and strength in the population. Our findings add to a growing body of evidence indicating mate choice behaviour can be a plastic trait, an idea that should be more explicitly considered in empirical studies of mate choice as well as conceptual frameworks of mate choice evolution and adaptive divergence.     

Consequences of elevated temperatures on life-history traits of an introduced fish, pumpkinseed Lepomis gibbosus

Life-history reactions of a pumpkinseed Lepomis gibbosus population in north-eastern France exposed to heated waters were studied. The study was conducted from 2001 to 2003 in an artificial reservoir, adjacent to a nuclear power plant, in which water temperatures are cool in winter (8·2–12·4° C) and rise early in spring (April: 14·7° C) nearly 5° C and 3° C over the temperature of its tributary, respectively. Fast growth among young-of-the-year, precocious maturity and short life span were observed, in contrast to related studies. The short life span appeared to be the price paid for early maturity in breeding fish, which suffered high mortality rates just after their first reproduction.     

The effect of elevated temperature on spawning of introduced pumpkinseed Lepomis gibbosus in Europe

In an experimental comparison of pumpkinseed Lepomis gibbosus reproduction under ambient and climate change water temperature regimes, spawning occurred earlier in the season, which is likely to lead to greater young-of-the-year survival with concomitant implications in the U.K. under warmer climatic conditions.     

Nest association of pumpkinseed, Lepomis gibbosus, and golden shiner, Notemigonus crysoleucas

I investigated the nest association of pumpkinseeds and golden shiners in an upstate New York pond. Golden shiners spawned in about one-third of pumpkinseed nests. Field observations indicate that golden shiners preferred to spawn in nests of male pumpkinseeds that attracted conspecific females, and avoided nests of pumpkinseeds that failed to do so. Golden shiners did not spawn in any of the nests that I kept clean experimentally after abandonment by male pumpkinseeds, suggesting that a clean nest without a guarding pumpkinseed is not enough to attract shiners to spawn. In field experiments, shiner eggs that were placed away from pumpkinseed nests suffered significantly higher losses to predation than those in the nests. This indicates that golden shiners benefit from spawning in pumpkinseed nests through the protection of their young by the host pumpkinseed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Variation in resource abundance affects diet and feeding morphology in the pumpkinseed sunfish (Lepomis gibbosus)

Summary Growth responses and accumulation of N and P were studied in two pygmy south-west Australian species of Drosera following supplementary feeding of arthropods (collembolans, Hypogastrura vernalis and fruit flies, Drosophila melanogaster) and/or a balanced mineral nutrient supplement (N as nitrate) via the roots. One feeding experiment used glasshouse-raised germlings from vegetative propagules (gemmae) of the perennial Drosera closterostigma, the other three (two on D. closterostigma and one on the annual D. glanduligera) involved natural populations engaging in natural captures of indigenous prey. All experiments recorded highly significant increases in plant dry matter, N and P (all plant age groups) and in reproductive performance (adult plants only) from artificial feeding of arthropods, but no apparent benefits from minerals alone or additive effects of minerals above that due to insects. Unresponsiveness to mineral nutrients was suggested to relate to inability of the species to use nitrate, while up to three-fold growth and nutrient uptake response to insects indicated that growth of natural populations might be severely limited by inadequate catches of prey. It is concluded that the highly nutrient-poor conditions typical of the habitat of pygmy species of Drosera may have promoted marked specialization towards carnivory and an attendant decline in ability to utilize soil-derived sources of nutrients.     

Genetic variation and phenotypic plasticity in a trophically polymorphic population of pumpkinseed sunfish (Lepomis gibbosus)

Summary Adaptive variation can exist at a variety of scales in biological systems, including among species, among local populations of a single species and among individuals within a single population. Trophic or resource polymorphisms in fishes are a good example of the lowest level of this hierarchy. In lakes without bluegill sunfish (Lepomis macrochirus), pumpkinseed sunfish (Lepomis gibbosus) can be trophically polymorphic, including a planktivorous limnetic form found in the pelagic habitat, in addition to the usual benthic form found in the littoral zone. In this paper we examine the degree to which morphological differences between the two forms are caused by genetic differences versus phenotypic plasticity. Adults from pelagic and littoral sites in Paradox Lake, NY, were bred separately and their progeny were raised in cages both in the open water and shallow water habitats of an artificial pond. The experimental design permitted two tests of genetic differences between the breeding stocks (in open and shallow water cages, respectively) and two tests of phenotypic plasticity (in the limnetic and benthic offspring, respectively). Limnetic progeny were more fusiform than benthic progeny raised in the same habitat. In addition, progeny of both stocks displayed limnetic-type characteristics when raised in the open water and benthic-type characteristics in the shallow water. Thus, genetic differences and phenotypic plasticity both contributed to the trophic polymorphism. Phenotypic plasticity and genetic differentiation accounted for 53 and 14%, respectively, of the variation in morphology. This study addresses the nature of subtle phenotypic differences among individuals from a single population that is embedded within a complex community, a condition that is likely to be the norm for most natural populations, as opposed to very large differences that have evolved in relatively few populations that reside in species-poor environments.     

Effects of predator exposure on baseline and stress-induced glucocorticoid hormone concentrations in pumpkinseed Lepomis gibbosus

We compared baseline and maximal cortisol concentrations between predator exposure and prey blood samples in pumpkinseed Lepomis gibbosus, captured using a standardised fishing event underneath osprey Pandion haliaetus nests and away from osprey nests. We did not detect differences in cortisol or glucose between sites. These findings suggest that predictable sources of predation risk may not confer stress-related costs in teleosts.     

Parasite fauna of introduced pumpkinseed fish Lepomis gibbosus: first British record of Onchocleidus dispar (Monogenea)

A survey of pumpkinseed Lepomis gibbosus from a stillwater fishery in southern England revealed infections by the ancyrocephalid monogenean Onchocleidus dispar at 100% prevalence. Host specificity of O. dispar to North American centrarchid fish suggests that it is a non-native parasite, introduced to Britain with L. gibbosus. Mean intensity of O. dispar was significantly higher in male (25.5 parasites) compared to female (15) and immature (7) hosts, but was not influenced by host standard length or sampling time. This sex-biased parasitism is likely to be related to both ecological and physiological factors, such as differential exposure to infective stages during nest building, or higher susceptibility to infection due to enhanced host stress levels. O. dispar significantly dominated the external surfaces and the outer gill arches. Analysis of the spatial distribution of the parasite between the 9 regions of gill surface determined that as density increased, higher numbers of parasites were found on the anterior and central gill regions compared to posterior (distal) regions. Apparent microhabitat selection is discussed in terms O. dispar life cycle, maximising oxygen availability, avoiding unstable areas and increasing intraspecific contact. The absence of O. dispar in the gills of roach, rudd and gudgeon sampled from the same fishery supports the assumption that this parasite is currently of little threat to native fish populations.     

Karyotype analysis of the pumpkinseed Lepomis gibbosus (Actinopterygii,Centrarchidae) by chromosomal banding and in situ hybridization

Sunfish are widely distributed in their native North American freshwaters and in many other geographic regions, including Europe. In this work the cytogenetics of L. gibbosus were studied. In particular, the authors localized the heterochromatic regions and the major and minor ribosomal gene families. The nucleolar organizer region was localized terminally in the short arm of only one pair, a large acrocentric pair, using both FISH and silver staining. Furthermore, the 5S ribosomal gene family was also localized by FISH in only one pair, in the centromeric region of the smallest chromosome pair. NOR and 5S rDNA regions were both C‐positive and CMA₃ positive. The CMA₃ positivity of the 5S ribosomal cluster is uncommon in fish, however, a similar situation has been found in M. salmoides, the only other centrarchid species studied with the same techniques. Moreover, the 5S ribosomal gene was sequenced and its molecular structure analysed.