Food of introduced mosquitofish: ontogenetic diet shift and prey selection

The mosquitofish Gambusia holbrooki of Lake Banyoles, Catalonia, Spain, were strictly littoral. Their population was dominated by females throughout the year, except in spring when males were more common. Their diet was based on littoral cladocerans, particularly Chydorus sphaericus, Scapholeberis ramneri, Ceriodaphnia reticulata , and Pleuroxus laevis , and nematoceran (basically chironomid) adults. There was a large variety of prey of terrestrial (collembolans, ants) or aquatic neustonic origin ( S. ramneri , emerging nematoceran adults), showing the microhabitat of mosquitofish closely linked to the water surface. In contrast to other studies, terrestrial insects were not especially important in summer. In addition to seasonal and between-site variation, there was an ontogenetic diet shift from microcrustaceans, particularly cladocerans (smallest fish also using diatoms and copepod nauplii) to larger prey, namely nematoceran adults. However, there was an unusual, inverse relationship between fish length and the size of two prey ( S. ramneri and the collembolan Sminthurides sp.), paralleling the overall increase in mean prey size.     

Ontogenetic diet shifts and interrupted piscivoryin introduced largemouth bass (Micropterus salmoides)

The ecology of largemouth bass (Micropterus salmoides) is one of the best known for freshwater fish, though largely through studies within its native range (North America). I studied the habitat and diet of a bass population introduced into a Mediterranean lake. The bass displayed strong ontogenetic diet shifts as follows: young‐of‐the‐year <25 mm fed on microcrustaceans; fish 25–75 mm, on amphipods and insects; fish 100–225 mm, on a freshwater shrimp, small fish and insects; fish 250–300 mm, on shrimp or crayfish; and fish >300 mm, on crayfish and large fish. The diet showed several differences from most previous studies: importance of freshwater shrimp instead of insects, low piscivory, and a delay in the ontogenetic shift to piscivory. Moreover, the ontogenetic shift to piscivory was interrupted at 250–300 mm, with consumption of shrimp and crayfish. This interruption of piscivory has been largely unreported and seems a consequence of the size‐structure and species composition of the fish assemblage. A review of the literature suggests that piscivory by largemouth bass might be generally lower in populations introduced outside North America.     

Survival benefits and divergence of predator-induced behavior between pumpkinseed sunfish ecomorphs

Resource use is widely thought to influence adaptive phenotypicdivergence, whereas other ecological factors, such as predation,are frequently overlooked, particularly in studies of polyphenismin fishes. Juvenile pumpkinseed sunfish (Lepomis gibbosus) rearedwith predatory walleye (Sander vitreus) increase body depthand dorsal spine length, indicating that developmental responsesto predation can shape phenotype. Body form responses to thesame predator cues though have also evolutionarily divergedbetween sunfish ecomorphs that coexist in single lake populationsby inhabiting either littoral or pelagic habitats, suggestingthat predation risk varies between habitats. Here, we test ifprior exposure to predator cues influences the development ofbehavior in juvenile pumpkinseed sunfish, if behavioral responsesto the same predator cues vary between ecomorphs, and if inducedphenotypic variation affects survival under predation. Behaviordepended strongly on prior exposure to predator cues, but thiseffect varied between sunfish ecomorphs, indicating that ecomorphshave different responses to the same predator cues. Predator-inducedphenotypes had higher survival than control phenotypes undersimulated littoral but not pelagic conditions. Predator-inducedphenotypic responses are candidate-inducible defenses, and divergentresponses between ecomorphs suggest that they can evolve inresponse to selection imposed by differences in habitat-specificpredation risk.     

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 environmental basis of diet variation in pumpkinseed sunfish, Lepomis gibbosus, and largemouth bass, Micropterus salmoides, along an Iberian river basin

The diet composition of largemouth bass, Micropterus salmoides, and pumpkinseed sunfish, Lepomis gibbosus, and its relation to environmental factors was studied in intermittent watercourses of the lower Guadiana basin (southern Iberia) during the particularly dry summer of 1994. Overall, both species took food items similar to those found in their North American and acclimatised ranges, with pumpkinseed consuming invertebrates (chiefly Chironomidae) and bass preying on invertebrates (chiefly Micronecta meridionalis) and fish (chiefly L. gibbosus and Gambusia holbrooki). Both bass and pumpkinseed were opportunistic feeders, feeding on the most frequent and abundant prey. However, Diptera larvae were apparently preferred by pumpkinseed and avoided by bass, while the contrary occurred with respect to Heteroptera. The two main fish prey of bass were eaten on the basis of random encounter. The relative abundance of macro-prey (i.e. fish and Atyephira desmarestii) were the principal environmental variables constraining bass dietary variation along the basin. On the contrary, pumpkinseed diet variation was mainly related to pumpkinseed size and the presence of piscivorous bass nearby, although habitat size and cyprinid abundance were also influential. This revised version was published online in July 2006 with corrections to the Cover Date.     

Contemporary and historical evolutionary processes interact to shape patterns of within-lake phenotypic divergences in polyphenic pumpkinseed sunfish, Lepomis gibbosus

Historical and contemporary evolutionary processes can both contribute to patterns of phenotypic variation among populations of a species. Recent studies are revealing how interactions between historical and contemporary processes better explain observed patterns of phenotypic divergence than either process alone. Here, we investigate the roles of evolutionary history and adaptation to current environmental conditions in structuring phenotypic variation among polyphenic populations of sunfish inhabiting 12 postglacial lakes in eastern North America. The pumpkinseed sunfish polyphenism includes sympatric ecomorphs specialized for littoral or pelagic lake habitats. First, we use population genetic methods to test the evolutionary independence of within-lake phenotypic divergences of ecomorphs and to describe patterns of genetic structure among lake populations that clustered into three geographical groupings. We then used multivariate analysis of covariance (MANCOVA) to partition body shape variation (quantified with geometric morphometrics) among the effects of evolutionary history (reflecting phenotypic variation among genetic clusters), the shared phenotypic response of all populations to alternate habitats within lakes (reflecting adaptation to contemporary conditions), and unique phenotypic responses to habitats within lakes nested within genetic clusters. All effects had a significant influence on body form, but the effects of history and the interaction between history and contemporary habitat were larger than contemporary processes in structuring phenotypic variation. This highlights how divergence can be better understood against a known backdrop of evolutionary history.     

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.     

Food of an endangered cyprinodont (Aphanius iberus): ontogenetic diet shift and prey electivity

Synopsis We studied the ontogenetic diet shift and prey electivity of an endangered cyprinodontid fish endemic to the Iberian Peninsula, the Spanish toothcarp (Aphanius iberus). The toothcarp’s diet was omnivorous, dominated by harpacticoid copepods (Mesochra lilljeborgi and Tisbe longicornis), copepod nauplii and detritus. Diet composition varied greatly among habitats, depending on prey availability. In a rarely inundated habitat (glasswort), there was more consumption of the isopod Protracheoniscus occidentalis and the harpacticoid copepod Mesochra lilljeborgi, while in algal mats another harpacticoid (Tisbe longicornis), chironomid dipterans and invertebrate eggs were more important in diet. Although a benthic feeding habitat has previously been suggested, in our study the diet was based rather on water column organisms for both glasswort and algal mat habitats. There was also an ontogenetic diet shift, with an increase of mean prey length with fish length, clearly linked to a microhabitat change. Smaller fish showed positive electivity and greater reliance on planktonic prey (e.g. copepod nauplii, the harpacticoid copepods Mesochra lilljeborgi and Tisbe longicornis, the rotifer Brachionus plicatilis, and ostracods), while larger fish elected and preyed on more benthic organisms (e.g. Canuella perplexa, Mesochra rapiens, and ephydrid dipterans).     

Apparent size choice of zooplankton by planktivorous sunfish: exceptions to the rule

Synopsis Choice of the apparently largest prey has been implicated as an important component of the size choice behavior of several planktivorous fish species. In this study we describe the effect of several aspects of prey placement, apparent and absolute size, and motion on the choice behavior of bluegill or white crappie. In binary choice experiments, bluegill usually choose Daphnia prey on the basis of apparent size. However, when both prey were close to the fish and the absolutely larger prey was apparently smaller, the fish commonly chose the absolutely larger. The horizontal placement of two prey also altered choice such that the more forward directed prey was chosen even when apparently smaller. White crappie, when offered a choice between a diaptomid copepod or daphnid prey, chose the daphnid most of the time. Bluegill sunfish offered moving versus non-moving heat-killed daphnids commonly chose the one in motion. Apparent size choice is still a good overall describer of bluegill and white crappie prey choice, but it is not the only mechanism involved in prey choice behavior of these fish.     

Differences in learning by foraging juvenile pumpkinseed and bluegill sunfish in a structured habitat

Synopsis We investigated the ability of two congeneric species of sunfish to learn to forage on a novel prey item in feeding arenas containing structured habitats. Eight bluegill sunfish and eight pumpkinseed sunfish were given the opportunity to forage on whiteworms daily for 10 days. Each day, several behavioural measures were recorded for each fish. Both species of sunfish learned to feed over the 10-day period but the bluegill sunfish learned to feed more quickly than the pumpkinseed sunfish. Pumpkinseeds, however, attained a higher level of foraging efficiency. The differences in learning and foraging efficiency were related to body morphology.     

Isolating the effects of ontogenetic niche shift on brain size development using pumpkinseed sunfish ecotypes

A functional relationship between relative brain size and cognitive performance has been hypothesized. However, the influence of ontogenetic niche shifts on cognitive performance is not well understood. Increases in body size can affect niche use but distinguishing nonecologically relevant brain development from effects associated with ecology is difficult. If survival is enhanced by functional changes in ecocognitive performance over ontogeny, then brain size development should track ontogenetic shifts in ecology. We control for nonecologically relevant brain size development by comparing brain growth between two ecotypes of Pumpkinseed sunfish whose ecologies diverge over ontogeny from a shared juvenile niche. Brain size differs between ecotypes from their birth year onwards even though their foraging ecology appears to diverge at age 3. This finding suggests that the eco‐cognitive requirements of adult niches shape early life brain growth more than the requirements of juvenile ecology.     

Ecological and morphological differentiation of pumpkinseed sunfish in lakes without bluegill sunfish

Summary Over the last three decades, sunfish of the familyCentrarchidae have become recognized as a model system in which the ecological consequences of species interactions can be observed and tested. The evolutionary consequences of species interactions in sunfish have received less attention. Bluegill (Lepomis machrochirus) and pumpkinseed (Lepomis gibbosus) sunfish are two common and well-studied species that occupy separate ecological niches. Adult bluegill are generalists that feed in the open water on zooplankton during much of the year, while adult pumpkinseeds specialize on crushing hard-bodied prey such as snails. These species coexist over much of their geographical ranges, but bluegill are historically absent from several large drainage basins in the northeastern US. Here we show that pumpkinseeds from an Adirondack lake without bluegills have differentiated into two morphological forms, one of which is planktivorous. Differentiation is independent of sex and occurs over a broad range of sizes. Thus, the ecological diversity that exists between the bluegill and pumpkinseeds in sympatry has been replaced by a comparable degree of diversity within pumpkinseeds in allopatry.     

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.     

Microhabitat preferences of longear sunfish: low light intensity versus submerged cover

Both cover and light intensity are important factors in microhabitat selection of fishes, suggesting a possible hierarchical relationship. Many centrarchids have been shown to prefer low light intensities and heavy submerged cover, but adult nonbreeding longear sunfish are reported to occupy sparsely vegetated, clear, shallow water during the brightest part of the day, probably for foraging. To determine the relative importance of cover and light intensity to longear sunfish, we presented nonbreeding adults with choices between these two conditions in a series of three laboratory experiments. Longear sunfish preferred low light to high light intensity and preferred submerged cover to no cover. However, when given a choice, they preferred to occupy low light intensity conditions without cover rather than submerged cover under higher light intensity conditions. Thus, low light intensity, such as is found in greater pool depths, may be the preferred refuge from predation for longear sunfish.     

Histological structure of the intestine and pyloric caeca of the green sunfish, Lepomis cyanellus Rafinesque

A histological study was made of the intestine and pyloric caeca of green sunfish, Lepomis cyanellus (Centrarchidae). The intestinal and caecal walls are histologically very similar, consisting of a mucosa (epithelial layer), submucosa (lamina propria and stratum compactum), muscularis (circular and longitudinal layers) and the serosa. Cellular constituents of the mucosal layer include absorptive, columnar epithelial cells, mucous-secreting goblet cells, and various leucocytes, the majority of which are lymphocytes. Other than relative size and the entrance of the bile duct at the base of the first caecum, no difference was found among caeca.
When fish were nutritionally stressed, a greater variety and number of leucocytes and shifts in the numbers of lymphocytes and goblet cells of the mucosal layer were the only observable effects.     

Origin and dispersal of bluegill sunfish, Lepomis macrochirus, in Japan and Korea

The bluegill sunfish, Lepomis macrochirus, is a notorious exotic species in many freshwater ecosystems, currently expanding its distribution worldwide. In 1960, a small group of bluegills captured in the Mississippi River at Guttenberg in Iowa were imported to Japan as a gift from the mayor of Chicago to the Japanese government. The offspring of these fish were released into the wild in Japan and also in Korea. Over 40 years after this first introduction, L. macrochirus now occupies all the freshwater ecosystems of both countries. We compared invading populations of L. macrochirus in Japan and Korea with native populations in the USA, using PCR-RFLP (polymerase chain reaction-restricted fragment length polymorphism) analyses of mitochondrial DNA, to estimate the origin and dispersal of L. macrochirus in Japan and Korea. Five haplotypes of mitochondrial DNA detected in Japanese and Korean populations completely coincided with the haplotypes of the Guttenberg population. Haplotype diversity of invading populations was shown to be highest in populations established in the 1960s, while genetic variability was lower in more recently established populations. Our results suggest that all L. macrochirus in Japan and Korea have originated from the 15 fish first introduced in 1960. Low haplotype diversity in newly established populations is probably due to genetic drift arising from repeated population bottlenecks, while the high similarity of haplotypes among neighbouring populations is considered to reflect the history of transplantation by humans.     

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.     

Age and size-dependent variation in the seasonal timing and probability of reproduction among mature female pumpkinseed,Lepomis gibbosus

Synopsis In this study, we used pumpkinseed,Lepomis gibbosus, populations in two east-central Ontario lakes to test for age and size-dependent effects on the probability and timing of reproduction of mature females. Pumpkinseed body size characteristics differed in the two lakes; Little Round Lake harbours a stunted population and Beloporine Lake does not. Age-class distribution of mature females was determined by a maturity assessment on fish collected just prior to spawning and at the mid-spawning period, combined with an early-season capture-mark-recapture survey. Spawning females were collected throughout the breeding season to assess age and size-related temporal trends, and to compare their age and size distribution with that of mature females at large. The proportion of age 2 females in Little Round Lake that actually spawned was considerably lower than the proportion of mature age 2 females at large. Furthermore, age 2 females that spawned in this lake did not do so until late in the breeding season. In contrast, the proportion of young/small females spawning in Beloporine Lake was comparable to the proportion of young/small mature females at large, and both small and large females spawned throughout the breeding season. Small mature females in Little Round Lake may have had no other option but to spawn late in the season because of their poor body condition. In Beloporine Lake, condition factor early in the breeding season in age 2–4 females was higher than that of Little Round Lake females, suggesting that limited energy reserves in the spring may have prevented young Little Round Lake females from early spawning. Our results show that the likelihood and timing of reproduction are both age and size-dependent in some populations. Small individuals that delay seasonal maturation and spawn late in the summer probably contribute little to the population due to the restricted growth and reduced overwinter survival of their progeny.To whom correspondence should be addressed     

New alien insect pests to Portugal on urban ornamental plants and additional data on recently introduced species

The spread of invasive pests is an important topic in pest control and early detection is crucial for containment of invasive aliens. In this study the results of recent surveys on alien insects of ornamental urban plants in Portugal are reported. Overall, 13 alien species associated with allochthonous ornamental plants were recorded. Seven of these pests are new to the Portuguese fauna: Tinocallis kahawaluokalani (Kirkaldy); Illinoia liriodendri (Monell); Appendiseta robiniae (Gillette) (Hemiptera: Aphididae); Acizzia jamatonica (Kuwayama) (Hemiptera: Psyllidae); Dasineura gleditchiae (Osten Sacken); Obolodiplosis robiniae (Haldeman) (Diptera: Cecidomyiidae); and Phytoliriomyza jacarandae Steyskal & Spencer (Diptera: Agromyzidae). Additional distribution data are also provided for other recently introduced alien pests in Portugal: Myzocallis (Lineomyzocallis) walshii (Monell) (Hemiptera: Aphididae); Platycorypha nigrivirga Burckhardt; Glycaspis brimblecombei Moore (Hemiptera: Psyllidae); Calophya schini Tuthill (Hemiptera: Calophyidae); Corythucha ciliata (Say) (Hemiptera: Tingidae); and Lantanophaga pusillidactylus (Walker) (Lepidoptera: Pterophoridae). Three species in particular (Illinoia liriodendri, Obolodiplosis robiniae and Phytoliriomyza jacarandae) were not previously found in the Iberian Peninsula. Details on current distribution, host plants and biological remarks are given for each species. Possible origin areas, pathways and impacts are also discussed.     

The role of propagule pressure in the invasion success of bluegill sunfish, Lepomis macrochirus, in Japan

The bluegill sunfish, Lepomis macrochirus, is a widespread exotic species in Japan that is considered to have originated from 15 fish introduced from Guttenberg, Iowa, in 1960. Here, the genetic and phenotypic traits of Japanese populations were examined, together with 11 native populations of the USA using 10 microsatellite markers and six meristic traits. Phylogenetic analysis reconfirmed a single origin of Japanese populations, among which populations established in the 1960s were genetically close to Guttenberg population, keeping high genetic diversity comparable to the ancestral population. In contrast, genetic diversity of later-established populations significantly declined with genetic divergence from the ancestral population. Among the 1960s established populations, that from Lake Biwa showed a significant isolation-by-distance pattern with surrounding populations in which genetic bottlenecks increased with geographical distance from Lake Biwa. Although phenotypic divergence among populations was recognized in both neutral and adaptive traits, P(ST)-F(ST) comparisons showed that it is independent of neutral genetic divergence. Divergent selection was suggested in some populations from reservoirs with unstable habitats, while stabilizing selection was dominant. Accordingly, many Japanese populations of L. macrochirus appear to have derived from Lake Biwa population, expanding their distribution with population bottlenecks. Despite low propagule pressure, the invasion success of L. macrochirus is probably because of its drastic population growth in Lake Biwa shortly after its introduction, together with artificial transplantations. It not only enabled the avoidance of a loss in genetic diversity but also formed a major gene pool that supported local adaptation with high phenotypic plasticity.