Dynamic intra‐epidermal bodies (IEBs) in koi epidermis

Koi scale epidermis contains large intra‐epidermal bodies (IEBs). IEBs are dynamic and circular structures that form in low frequency within the epidermis. During formation, an IEB pulls down the microridge‐laden surface layer, which takes on a creased or wrinkled appearance. After the IEB constricts, the microridge layer unfolds to its original state. The newly described IEBs are distinctly different from the recently reported apical rings which are situated on the surface of individual epidermal cells. While apical rings are directly exposed to the external environment where sampling can occur, IEBs appear to reflect intra‐epidermal events such as sequestering of dead cell remnants.     

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.     

Patterns of reproduction in two populations of pumpkinseed sunfish,Lepomis gibbosus,with differing food resources

Synopsis The influence of available food resources on growth and patterns of reproduction was studied in two populations of pumpkinseed sunfish in adjacent lakes in southeastern Ontario. Ideally, mature individuals should be long-lived, attain a large size, and maintain a high reproductive effort. However, rarely are environmental conditions ideal and compromises are necessary. In the present study, there were smaller quantities of preferred prey items (gastropods), and total benthos available in one lake. Pumpkinseed in this population ate less in general, and were smaller in body size in the older age classes. These fish matured at a younger age and a smaller size than individuals in the other lake studied, and in populations in this geographic area reported in the literature. In addition, the relationships of both fecundity and ovary weight to total weight were different between lakes. Mean ovary weight, ripe egg size and lifetime fecundity were less in the population of small-bodies fish. Variation in patterns of reproduction between populations suggests that fish may be able to fine-tune responses to local environmental conditions.     

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.     

Effects of increasing temperature on social behaviour in territorial groups of pumpkinseed sunfish,Lepomis gibbosus

To study the effect of thermal stress on their social behaviour, ten territorial groups of pumpkinseed sunfish, Lepomis gibbosus, were subjected to a 1°C temperature increase every other day until they succumbed. Social behaviour remained remarkably unchanged by thermal stress until nearly lethal levels. As temperature rose, ritualised behaviours increased in frequency, then fell off, probably reflecting changes in general activity.Behavioural and physiological signs of stress appeared from 31 to 38°C. Events during these social breakdowns are described and compared with the response of two other freshwater fishes with different social organisations subjected to the same thermal regime.     

Growth energy partitioning by juvenile bluegill sunfish, Lepomis macrochirus Rafinesque

Growth energy build-up and partitioning by immature bluegill sunfish were studied over the summer months from May to September 1984. Fish sampled in May showed low condition and low lipid reserves. Condition was restored to summer levels within the following month. Fat reserves were built up from 7% to 14% of body weight over the entire summer. Most (80%) of the annual length increase occurred from 1 June to 15 August. Over the winter period (October-April) condition dropped and lipids were depleted. Calculations showed that energy liberated through lipid depletion over this period could supply the energy requirements of inactive wintering fish. It is concluded that the sequencing of growth energy diversion into changes in body length, condition, and lipid build-up enhances mid-summer growth and overwinter survival.     

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.     

Interpopulation variation in growth and life-history traits of the introduced sunfish, pumpkinseed Lepomis gibbosus, in southern England

We examined attributes of growth and reproduction in 19 populations of pumpkinseed (Lepomis gibbosus) introduced into southern England in order to: (i) assess variability of these traits in a northern European climate; (ii) assess inter‐relationships among these variables; and (iii) compare these attributes with populations from other parts of Europe where pumpkinseeds have been introduced. Growth rates varied considerably among populations, but juvenile growth rates and adult body sizes were generally among the lowest in Europe. Mean age at maturity ranged from 2.0 to 3.9, and was strongly predicted by the juvenile growth rate (earlier maturity with faster juvenile growth). Other population parameters that also displayed significant negative associations with mean age at maturity were gonadosomatic index, body condition, and adult body size (total length, TL at age 5). Mean TL at maturity and the adult growth increment showed no significant associations with any of the other growth or life‐history variables. Pumpkinseed populations in England matured significantly later than those introduced into warmer, more southerly areas of the continental Europe. All of these data suggest that a combination of cool summer temperatures and resource limitation is the cause of slow growth, small adult body size and delayed maturity relative to introduced populations on the European mainland.     

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.     

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.     

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.     

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.     

Sperm investment and alternative mating tactics in bluegill sunfish (Lepomis macrochirus)

Although alternative mating tactics are found in males of manyspecies, little is known about tactic-specific adaptations tosperm competition and the mechanism by which fertilization successis obtained. We now report on the sperm investment patternsof males that use alternative mating tactics in bluegill sunfish(Lepomis macrochirus). Bluegill males are characterized by threealternative mating tactics: parentals, sneakers, and satellites.Parentals delay maturation and adopt a courting and guardingtactic, whereas sneakers and satellites mature precociouslyand, respectively, use sneaking and female mimicry to stealfertilizations from parentals. We found differences among thetactics in testes weight, sperm longevity, and ejaculate spermdensity and competitiveness. Parentals had the largest testes,but the smallest relative to their body weight. Satellites hadintermediate-sized testes, and sneakers had the smallest testes,but the largest relative to their body weight. Within each tactic,there was a positive relationship between testes weight andbody weight, but the exact relationship differed among the tacticsand could not be attributed solely to allometry. Parentals hadlonger-lived sperm compared with that of both sneakers and satellites.Ejaculate sperm density was greatest in sneakers, satelliteswere intermediate, and parentals had the lowest. Competitionexperiments involving equal volumes of ejaculate showed thatfertilization success increased linearly with sperm number,consistent with a rafflelike process. However, independent ofsperm number, parental sperm were superior at fertilizing eggs.We interpret these sperm investment patterns in relation todifferences in sperm competition risk, number of mating opportunities,and alternative investment options.     

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.     

Post-mating reproductive barriers in two unidirectionally hybridizing sunfish (Centrarchidae: Lepomis)

The evolutionary sequence of events in the evolution of reproductive barriers between species is at the core of speciation biology. Where premating barriers fail, post-mating barriers, such as conspecific sperm precedence (CSP), gamete incompatibility (GI) and hybrid inviability (HI) may evolve to prevent the production of (often) costly hybrid offspring with reduced fitness. We tested the role of post-mating mechanisms for the reproductive isolation between two sunfish species [bluegill (BG) Lepomis macrochirus and pumpkinseed (PS) Lepomis gibbosus] and their first-generation hybrids. Performing in vitro sperm competition experiments, we observed asymmetric CSP as main post-mating isolation mechanism when BG and PS sperm were competing for PS eggs, whereas when sperm from both species were competing for BG eggs it was HI. Furthermore, hybrid sperm--although fertile in the absence of competition--were outcompeted by sperm of either parental species. This result may at least partly explain previous observations that natural hybridization in the study system is unidirectional.     

Condition-dependent nesting in bluegill sunfish Lepomis macrochirus

1. There are two prominent, nonmutually exclusive hypotheses to explain the timing of reproduction in animals: energetic constraint and adaptive behaviour. 2. We tested these hypotheses by quantifying the costs and benefits of nesting at different times in the season for male bluegill sunfish Lepomis macrochirus, a species with paternal care, in Lake Opinicon (Ontario, Canada). 3. The value of nesting at different times during the breeding season (RSb) was determined from spawning individuals as RSb = Pb x Cb x Sb x Ob, where Pb is the probability of spawning during each bout b, Cb is the expected brood size, Sb is the expected brood survivorship to 'swim-up', and Ob is the survivorship of free-swimming fry to age 1 year. 4. The results show that the value of nesting peaks during the middle of the season. However, nesting patterns varied with male condition and not all males nested at the peak. 5. Larger males, which were able to nest multiple times, first nested early in the season when overwinter survivorship of offspring and renesting opportunities later in the season were maximized. These males had the highest seasonal reproductive success. 6. Smaller males, which nested a single time, delayed nesting until the middle of the season when spawning opportunities and brood sizes were greatest. 7. These data suggest that both energetics and adaptive behaviour play roles in determining the timing of reproduction.