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.     

Nontroglobitic Fishes in Bruffey-Hills Creek Cave, West Virginia, and other Caves Worldwide

Six species of fishes were tagged and released in September and November 1995 and on five dates between November 1998 and October 1999 inside Bruffey-Hills Creek Cave. Most of the tagged fishes were creek chub, Semotilus atromaculatus, and green sunfish, Lepomis cyanellus. The overall recapture rate was 2.6% as only three of 117 fishes were recaptured. Forty-nine days was the longest time of residence by two L. cyanellus and one S. atromaculatus, and movement of 83.4 m was observed only for the creek chub. Tag loss was confirmed as one factor in the low recapture rate. Nine species of fishes were collected in the cave, including Phoxinus oreas and Pimephales promelas, two species never collected in a cave. Most of the fishes were pigmented normally, but many creek chubs were depigmented and appeared white or pigmentless when first observed in the cave stream. Four species, Phoxinus oreas, Pimephales promelas, Ameiurus nebulosus, and Lepomis macrochirus, were collected for the first time in Bruffey-Hills Creek Cave and in the Bruffey and Hills creeks drainage. Although data from this study did not shed light on residence time of fishes in the cave, the consistent occurrence of epigean fishes in this cave was shown. A list of epigean fishes from caves worldwide was included along with a discussion of aspects of the ecology of epigean fishes in caves and of evolution of troglobitic fishes.     

Implications of stunting on morphology of freshwater fishes

The purpose of this study was to assess morphological differences between stunted and non‐stunted white perch Morone americana and green sunfish Lepomis cyanellus. Few female M. americana were captured; thus, morphological differences between adult males and juveniles were assessed for M. americana. Similarly, few immature (juvenile) L. cyanellus were captured for the stunted morphotype; thus, male and female morphological differences were assessed for L. cyanellus. Features of the head tended to be relatively larger in stunted fish of both species, whereas the mid‐body tended to be relatively larger in non‐stunted M. americana, but not in non‐stunted L. cyanellus. Adult and juvenile morphology overlapped considerably in non‐stunted M. americana, but there was a clear distinction between adult and juvenile morphology of stunted M. americana. There was little sexual dimorphism in shape in stunted L. cyanellus, whereas sexual dimorphism was evident in non‐stunted L. cyanellus. It appears that selective forces imposed by predation and food limitation may contribute to morphological diversification between stunted and non‐stunted fishes.     

Variation in the diet and feeding morphology of polyphenic Lepomis macrochirus

Bluegill Lepomis macrochirus showed variation in their diet and trophic morphology based on habitat. Pelagic L. macrochirus feed almost exclusively on cladocerans; littoral L. macrochirus feed on a variety of benthic invertebrates, molluscs, cladocerans and insects. Fish from the littoral habitat had wider pharyngeal jaws, which probably allowed them to crush gastropods and process benthic invertebrates.     

Getting in shape: habitat‐based morphological divergence for two sympatric fishes

Freshwater fishes often show large amounts of body shape variation across divergent habitats and, in most cases, the observed differences have been attributed to the environmental pressures of living in lentic or lotic habitats. Previous studies have suggested a distinct set characters and morphological features for species occupying each habitat under the steady–unsteady swimming performance model. We tested this model and assessed body shape variation using geometric morphometrics for two widespread fishes, Goodea atripinnis (Goodeidae) and Chirostoma jordani (Atherinopsidae), inhabiting lentic and lotic habitats across the Mesa Central of Mexico. These species were previously shown to display little genetic variation across their respective ranges. Our body shape analyses reveal morphometric differentiation along the same axes for both species in each habitat. Both possess a deeper body shape in lentic habitats and a more streamlined body in lotic habitats, although the degree of divergence between habitats was less for C. jordani. Differences in the position of the mouth differed between habitats as well, with both species possessing a more superior mouth in lentic habitats. These recovered patterns are generally consistent with the steady–unsteady swimming model and highlight the significance of environmental forces in driving parallel body shape differences of organisms in divergent habitats. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 152–162.     

Fish mortality and physicochemistry in a managed floodplain wetland

Patterns of fish mortality and associated physicochemical factors werestudied during late spring in a managed wetland canal along the lowerMissouri River, Missouri. Mean dawn dissolved oxygen was lower and meanun-ionized ammonia and turbidity were higher during the fish kill thanbefore or after the kill, or than was observed in a nearby wetland canalwhere no fish kill occurred. Dissolved oxygen at dawn and un-ionizedammonia concentrations were at critically low and high levels respectively,so that both likely contributed to the fish mortality. Timing and magnitudeof observed carcasses suggested that Ameiurus melas Rafinesques wasthe most tolerant species for the sizes observed compared to Ictiobuscyprinellus Valenciennes, Lepomis macrochirus Rafinesque, Cyprinus carpio Linneaus, and Lepomis cyanellus Rafinesque.Decreasing mean lengths of fish carcasses during the fish kill for C.carpio, L. cyanellus, and A. melas, indicate that smaller fishes mayhave been more tolerant of harsh environmental conditions than largerindividuals of the same species. Differential mortalities among species andsizes during drawdowns in actively managed wetland pools may haveintentional and unintentional ramifications on wetland and riverine fishcommunity structure, fish-avian interactions, and implementing anecosystem management perspective to restoring more naturalized riverfloodplain wetland functions. Late summer and early autumn draining ofmanaged wetlands might be used to benefit a wider diversity of wildlife andfishes.     

Utilizing stomach content and faecal DNA analysis techniques to assess the feeding behaviour of largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus

In this study, the feeding behaviour of the non‐native invasive predatory fishes largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus was studied in the Ezura River, a northern tributary of Lake Biwa, Japan. Prey composition was estimated based on visual examination of stomach contents and faecal DNA analysis to determine feeding habits of these predatory fishes. Stomach content analysis showed that native fishes (e.g. ayu Plecoglossus altivelis and gobies Rhinogobius spp.) and shrimps (e.g. Palaemon paucidens) were the major prey items for M. salmoides, while snails, larval Chironomidae and submerged macrophytes were the dominant prey items of L. macrochirus. Micropterus salmoides tended to select larger fish in the case of crucian carp Carassius spp., but smaller fishes in the case of P. altivelis and Rhinogobius spp. Faecal DNA analyses revealed prey compositions similar to those identified in predator stomach contents, and identified additional prey species not detected in stomach content inspection. This study demonstrated that both stomach content inspection and DNA‐based analysis bear several inherent shortcomings and advantages. The former method is straightforward, although identification of species can be inaccurate or impossible, whereas the latter method allows for accurate species identification, but cannot distinguish prey size or stage. Hence, integration of morphology‐based and DNA‐based methods can provide more reliable estimates of foraging habits of predatory fishes.     

Elevated carbon dioxide has limited acute effects on Lepomis macrochirus behaviour

The current study investigated the behavioural response of Lepomis macrochirus following exposures to elevated carbon dioxide (CO₂). For this, L. macrochirus were held at ambient pCO₂ (160 μatm pCO₂) for 7 days, then exposed to elevated pCO₂ (8300 μatm pCO₂) for 5 days, and then returned to ambient conditions for a further 5 days to recover. At the end of each exposure period, several behavioural metrics were quantified (boldness, lateralization and activity). Data showed no change in lateralization and most metrics associated with performance and boldness. During the boldness test, however, average velocity, velocity in the thigmotaxis (outer) zone and proportion of activity in the thigmotaxis zone increased with pCO₂ exposure. During post‐exposure, average velocity of L. macrochirus decreased. In addition, individual rank was repeatable during the pre‐exposure and post‐exposure period in three of the 17 metrics investigated (average velocity in the middle zone, average velocity near object and total shuttles to the object zone), but not during the CO₂ exposure period, suggesting that elevated pCO₂ disrupted some behavioural performances. Overall, this study found elevated pCO₂ caused disruption to behaviours of freshwater fishes such as L. macrochirus and effects do not appear to be as serious as has been shown for marine fishes.     

Planktivory by bluegill (Lepomis macrochirus) on Leptodora kindti in a small North American lake

Leptodora kindti (Crustacea: Cladocera) is a large species of zooplankton (2–18 mm length) that is exceptionally transparent. This transparency is believed to be a means by which it successfully coexists in lakes with planktivorous fishes. We investigated the gut remains of bluegill (Lepomis macrochirus) that had been feeding on L. kindti and Daphnia (D. galeata and D. retrocurva) in the wild (Lake Zurich, Illinois) and found that bluegill readily preyed on L. kindti as small as 3–5 mm length, and strongly selected L. kindti over Daphnia galeata and Daphnia retrocurva. The large compound eye of L. kindti is one half to one complete order of magnitude larger than Daphnia's eye, consistent with the hypothesis that eye area is an important visual cue for fishes. Moreover, the slope of the relationship between eye area and body length is an order of magnitude shallower in L. kindti than Daphnia, suggesting that eye area has been under stronger negative selection in L. kindti. Results suggest that L. kindti's large and dark eye compromises the transparent nature of its body.     

Nest‐site fidelity in parental male bluegill Lepomis macrochirus: spatial patterns and the influence of prior mating success

A 4 year mark–recapture study examined the pattern of nesting site fidelity of parental‐type male bluegill Lepomis macrochirus. The study results indicated that iteroparous male L. macrochirus choose new nest sites near their own previously used sites. The scale of site fidelity varied, but generally males choose to renest within shoreline areas rather than specific or exact nest locations (94% within‐year, 86% among‐years). Iteroparous males also displayed no preference to nest in proximity to neighbouring males from previous colonies to suggest social fidelity. Contrary to expectation, manipulating males' reproductive success had no significant effect on the pattern or scale of male reproductive site fidelity.     

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.     

Group swimming behaviour and energetics in bluegill Lepomis macrochirus and rainbow trout Oncorhynchus mykiss

Group swimming size influences metabolic energy consumption and swimming behaviour in fishes. Hydrodynamic flows and vortices of other fish are thought to be beneficial in terms of the energetic costs of swimming. Similarly, abiotic obstructions have been shown to have similar benefits with respect to metabolic consumption in swimming fish such as rainbow trout Oncorhynchus mykiss. The current study works to examine metabolic rates and swimming behaviours as a function of group swimming with bluegill sunfish Lepomis macrochirus and O. mykiss. Fishes were subjected to individual and group swimming in a respiratory swim tunnel to determine oxygen consumption as a proxy for the metabolic rate of swimming fish. In addition, fish movements within the swim tunnel test chamber were tracked to examine group swimming behaviour. We hypothesized that fish would benefit metabolically from group swimming. In the case of O. mykiss, we also hypothesized that groups would benefit from the presence of an abiotic structure, as has been previously observed in fish swimming individually. Our results suggest that the influence of group size on swimming metabolism is species specific. While L. macrochirus show decreased metabolic rate when swimming in a group compared to individually, O. mykiss did not show such a metabolic benefit from group swimming.     

Social familiarity and shoal formation in juvenile fishes

The potential influence of social familiarity in shoal‐choice decisions was investigated in two sympatric species of north temperate fishes, juvenile banded killifish Fundulus diaphanus and juvenile bluegill sunfish Lepomis macrochirus. Groups of socially familiar and unfamiliar conspecifics were formed in the laboratory using wild‐caught fishes. Juvenile F. diaphanus demonstrated a strong preference for familiar conspecific shoalmates, whereas juvenile L. macrochirus exhibited no preference for either unfamiliar or familiar conspecific shoalmates. The differential influence of familiarity on shoalmate choice in juveniles of these two species could be due to their different ecologies, local population densities and life histories.     

Otolith chemistry of prey fish consumed by a fish predator: does digestion hinder Russian doll techniques?

The effect of digestion by a predatory fish (largemouth bass Micropterus salmoides) on stable isotopic (δ¹³C and δ¹⁸O) and trace elemental (Sr:Ca and Ba:Ca) compositions of prey fish (bluegill Lepomis macrochirus) otoliths was investigated in a laboratory experiment. Trace element and stable‐isotopic signatures of L. macrochirus otoliths were not significantly altered for up to 16 h after L. macrochirus were consumed by M. salmoides. Prey fish otoliths recovered from predator digesta can retain environmental stable isotopic and trace elemental signatures, suggesting that determination of environmental history for prey fishes by stable‐isotope and trace‐element analysis of otoliths recovered from stomachs of piscivorous fishes will be feasible.     

Utilization of autochthonous macroinvertebrate drift by a pool fish community in a woodland stream

Qualitative and quantitative drift data were collected simultaneously above and below a pool both before and after the pool was heavily electrofished. These data revealed no significant difference between drift organism densities (#/m³) above or below the pool before or after fish collection. Qualitative and quantitative fish gut analyses suggest that the sunfishes Lepomis megalotis, L. macrochirus and L. cyanellus may be feeding on drifting invertebrates as these organisms comprised 58%, 37% and 35% gut volume, respectively. A design for a long-term drift net apparatus is presented.     

Functional morphology of the tetra fish Astyanax lacustris differs between divergent habitats in the Pantanal wetlands

This study investigated whether the body morphology of the tetra fish Astyanax lacustris (previously Astyanax asuncionensis) varied between populations inhabiting one lagoon (a lentic, shallow environment, with great habitat complexity created by aquatic macrophytes) and an adjacent river (a deeper, lotic environment where aquatic macrophytes are scarce) in a seasonally flooded wetland, despite population mixing during the wet season. Morphological differences matched a priori predictions of the theory relating functional body morphology and swimming performance in fishes between lagoon and river habitats. Observed morphological variation could have resulted from adaptive habitat choice by tetras, predation by piscivores and adaptive phenotypic plasticity during development.     

Maintenance of female mimicry as a reproductive strategy in bluegill sunfish (Lepomis macrochirus)

Synopsis In species where male reproductive success is dependent on male competition and aggression, alternative reproductive patterns, thought to represent a reduction in male reproductive effort, sometimes occur. Female mimicry in bluegill sunfish (Lepomis macrochirus) is an example of an obligate alternative male strategy. Female mimics are small, sexually mature males which mimic the details of female behavior, and gain access to functional females attracted to the nests of large, aggressive territorial males. The costs of female mimicry relative to nesting male behavior are discussed and two hypotheses, deception and mutual gain, are presented to explain the tolerance and courtship of female mimics by nesting males.     

Diet of bluegill Lepomis macrochirus in a South African reservoir during winter and summer

Alien fishes are considered a major threat to aquatic biodiversity in South Africa, yet relatively little regional information on their biology and ecology is available for many of these species. Seasonal changes in the diet of the bluegill Lepomis macrochirus in Howieson’s Poort Dam, Grahamstown, were assessed during summer and winter in 2014–2015, using stomach content analysis. In winter, juvenile and adult fish diets were dominated by crustacean zooplankton and insects, respectively. In summer, juvenile fish fed on crustaceans and insects, whereas adults consumed mostly fish eggs, indicating a potential impact by these invasive fish on native fish through oophagy.     

The use of floating overhead cover by warmwater stream fishes

Increasing the amount of woody debris in streams has often increased population size of one or more focal species whereas clearing a stream of woody debris has often reduced populations. Alterations of the amount of woody debris change multiple aspects of the habitat simultaneously, so it is very difficult to know what particular stimulus or combination of stimuli evoked the changes in the fish populations. The purpose of this research was to alter habitat by the addition of overhead cover alone, and to see whether or not that single change would affect the distribution of stream-dwelling fishes. The addition of solid, floating cover objects 1.8 m by 2.4 m resulted in increased local populations of bluntnose minnow (Pimphales notatus), creek chub (Semotilus atromaculatus), and longear sunfish (Lepomis megalotis) five to six weeks later, but did not change the distribution of four other species: white sucker (Catostomus commersoni), green sunfish (Lepomis cyanellus), bluegill (Lepomis macrochirus), and largemouth bass (Micropterus salmoides). These results showed that for some species at least, changes in channel morphology, visual isolation, and amelioration of flow produced by adding three-dimensional complex woody debris were not essential for making locally attractive habitat changes in a warmwater stream. The effectiveness of overhead cover as used here would be expected to vary for different species and depend on such things as predation threat, flow, and food levels.     

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.