Trophic polymorphism in introduced pumpkinseed (<Emphasis Type="Italic">Lepomis gibbosus</Emphasis>) inhabiting Iberian reservoirs

Introduced pumpkinseed in Iberian reservoirs display marked external morphological differentiation along two simultaneous dimensions of flow and trophic structure. We assessed the degree of internal morphological differentiation using gill rakers, pharyngeal jaws and the levator posterior muscle among pumpkinseeds occupying four different habitats and determined whether prey consumption accounted for any discernible differences in feeding structures among ecomorphs. Results showed significant differentiation by habitat based on pharyngeal muscle and jaw dimensions in all study reservoirs, with pelagic pumpkinseeds having smaller jaws than littoral pumpkinseeds in four of the five reservoirs. Gill rakers, however, differentiated morphs in only three of the five reservoirs, corresponding to differences in zooplankton consumption among pelagic and littoral individuals in those reservoirs. Based on all internal morphological traits, greater divergence was seen along the littoral-pelagic trophic axis in the lacustrine zones of reservoirs compared to the fluvial zone. Overall differences noted in internal morphology are likely the result of phenotypic plasticity; the ability of this species to readily adapt to changing physical environments may explain the success of the pumpkinseed in its introduced range.     

Morphological and life-history differentiation between littoral and pelagic forms of pumpkinseed

Trophically dimorphic pumpkinseed populations were investigated in five lakes in Ontario, Canada to determine: (1) whether the morphological traits that distinguish littoral from pelagic forms are consistent among populations; (2) whether the pharyngeal jaw apparatus is diminished in pelagic pumpkinseeds because of a lower proportion of hard-bodied prey in their diets; (3) whether there is life-history differentiation between littoral and pelagic forms. Pumpkinseeds captured from the littoral and pelagic zones differed significantly in morphology in four of the five lakes, but the number of external measures that differed significantly within the differentiated populations ranged from zero to six. Littoral pumpkinseeds generally had longer heads, more rearward placement of dorsal and pectoral fins, longer pectoral fins and deeper bodies than pelagic pumpkinseeds. Littoral and pelagic pumpkinseeds were more readily differentiated by internal morphometric measures, with littoral individuals having larger molars and wider spacing between gill rakers than pelagic individuals. Littoral and pelagic differences in age at maturity, size at maturity and gonado-somatic index were present only in one of three populations assessed for these traits, suggesting that morphological divergence is not necessarily accompanied by life-history differentiation.     

Analyzing nested variation in the body form of Lepomid sunfishes

Phylogenetic hierarchies are often composed of younger diverging lineages nested within older diverging lineages. Comparing phenotypic variation among several hierarchical levels can be used to test hypotheses about selection, phenotypic evolution and speciation. Such hierarchical comparisons have only been performed in threespine stickleback, and so here we use a hierarchical pattern of divergences between near-shore littoral and off-shore pelagic habitats to test for selection on the evolution of body form in Lepomis sunfish in lakes. We compare variation in external body form between fish from littoral and pelagic habitats at three levels: among ecomorphs within individual lake populations (intrapopulation), among populations of the same species in different lakes (interpopulation), and between bluegill and pumpkinseed sunfish species (interspecifically). Using geometric morphometric methods, we first demonstrate that interpopulation variation in mean body form of pumpkinseed sunfish varies with the presence of pelagic habitat. We then incorporate these results with existing data in order to test the similarity of phenotypic divergence between littoral and pelagic habitats at different hierarchical levels. Parallel relationships between certain body form traits (head length, caudal length and pectoral length) and habitat occur at all three levels suggesting that selection persistently acts at all levels to diversify these traits and so may contribute to species formation. For other traits (caudal depth and pectoral altitude), divergence between habitats is inconsistent at different hierarchical levels. Thus, nested biological variation in Lepomid body form reflects a history of deterministic selection and historical contingency, and also identifies traits that likely have likely influenced fitness and so serve important functions.     

Asymmetrical habitat coupling of an aquatic predator—The importance of individual specialization

Predators should stabilize food webs because they can move between spatially separate habitats. However, predators adapted to forage on local resources may have a reduced ability to couple habitats. Here, we show clear asymmetry in the ability to couple habitats by Eurasian perch—a common polymorphic predator in European lakes. We sampled perch from two spatially separate habitats—pelagic and littoral zones—in Lake Erken, Sweden. Littoral perch showed stronger individual specialization, but they also used resources from the pelagic zone, indicating their ability to couple habitats. In contrast, pelagic perch showed weaker individual specialization but near complete reliance on pelagic resources, indicating their preference to one habitat. This asymmetry in the habitat coupling ability of perch challenges the expectation that, in general, predators should stabilize spatially separated food webs. Our results suggest that habitat coupling might be constrained by morphological adaptations, which in this case were not related to genetic differentiation but were more likely related to differences in individual specialization.     

Morphological differences between native and non-native pumpkinseed in traits associated with locomotion

Adaptive strategies in morphology can significantly influence the successful invasion and establishment of non-native species. Since its introduction, the pumpkinseed (Lepomis gibbosus), a sunfish of North American origin, has spread throughout most of Europe, including the Iberian Peninsula. We hypothesized that 12 morphological traits, functionally significant for locomotion, would differ according to geographic origin (native/non-native) and habitat type (fluvial/lacustrine). Using flow-through raceways, we simultaneously reared F₁ young-of-the-year pumpkinseed from two native and two non-native populations, produced from adults kept in a common environment. Morphometric measurements were recorded at the beginning and end of the 90-day rearing period. Median-fin size and placement differed significantly between native and non-native populations, whereas paired fin size differed between fluvial and lacustrine populations. Other functionally significant traits, such as body width, also differed between native and non-native populations. Spanish populations were considered to have acquired these adaptive external morphologies through successive generations, following the species’ range expansion through the variable environments of the Iberian Peninsula.     

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.     

Changes in use of lake habitat by experimentally segregated populations of cutthroat trout and Dolly Varden char

The role of competition in a lacustrine community of two salmonid species, cutthroat trout Oncorhynchus clarki and Dolly Varden char Salvelinus malma , was studied in three coastal British Columbia lakes Habitat use by the species alone (allopatric) and in coexistence with each other (sympatric) was investigated by gill netting at 0-40 m depth contours (surface to bottom) so that several habitats (littoral, epipelagic. pelagic, epibenthic) were sampled From June to October, trout used mainly littoral and epipelagic habitats in sympatry and allopatry Char used all habitats in allopatry. and deep habitats (pelagic, epibenthic) not frequented by trout in sympatry The two species were thus spatially segregated with depth in sympatry Diel (day, night) and seasonal changes (spring, summer, autumn) in habitat use were not pronounced The shift in habitat use by experimentally allopatric char but not trout suggests that the effects of competition between sympatric trout and char for habitat resources are greater on the char     

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.     

Effects of habitat and food resources on morphology and ontogenetic growth trajectories in perch

Studies on resource polymorphism have mainly been considered at the end stage of ontogeny, whereas many species undergo diet changes as they grow. We conducted a field survey to analyze the role of adaptive variation during ontogeny in Eurasian perch (Perca fluviatilis). We caught perch from the littoral and pelagic zones of a lake to investigate whether perch differ in morphology and diet between these habitats. We also investigated whether there were any differences in morphological trajectories during the ontogeny of perch collected from the two habitats. We found that perch caught in the littoral habitat, independently of size, had a deeper body, larger head and mouth and longer fins than perch caught in the pelagic zone. Macroinvertebrates and fish dominated the diet of littoral perch, whereas the diet of the pelagic perch consisted mainly of zooplankton and to some extent fish. Independently of size, the more streamlined individuals had a larger proportion of zooplankton and a smaller proportion of macroinvertebrates in their diet than the deeper-bodied individuals, indicating a relation between diet and morphology. Some morphological characters followed different ontogenetic trajectories in the two habitats; e.g. the changes to a deeper body and a larger head were faster in the littoral than in the pelagic perch. The relationship between the length of perch and the size of the mouth and fins also differed between perch from the two habitats, where the increase in the length of the pelvic fin and the area of the mouth increased faster with size in the littoral perch. Our findings show that variation in morphology between habitats differs during ontogeny in a way that corresponds to functional expectations for fish species that occupy these habitats.     

Fish in Reservoirs

In all reservoirs the fish fauna is recruited from their riverine antecedents. The nature of the riverine environment, created by its unstable hydrological regime, favours the evolution of generalist species. Of these species, only a few are preadapted to lacustrine conditions. When riverine fish communities are trapped in a reservoir, most of the fish stay close to the shore, the mouth of tributaries and in shallows. The pelagic and deep waters are poorly utilized and the yield of fish is below the natural productivity of these water bodies, especially in the case of deep and large reservoirs. This occurs unless the dammed river contains fish species from families which are preadapted for lacustrine conditions. A positive correlation between the percentage of pelagic fish and the total yield of fish in reservoirs indicates that the pelagial represents a vacant habitat, only seldom invaded by riverine species. This situation probably applies to the deep water of reservoirs too, although sufficient data to demonstrate this are not available. The littoral region of reservoirs is utilized by some generalists of euryoecious species derived from the indigenous riverine fauna which may be preadapted for lacustrine conditions. If such preadapted lacustrine species are not present, the introduction of lacustrine species or those preadapted for the lacustrine environment is desirable under certain circumstances. However, caution must be exercised in any introduction.     

高营养级捕食者在浅水湖泊沿岸带与敞水区能量耦合的维持作用

研究利用保安湖沿岸带与敞水区两种生境中高营养级捕食者(达氏鲌与红鳍原鲌)的碳(δ13C)、氮(δ15N)稳定同位素数据,通过稳定同位素质量平衡混合模型、非度量多维尺度分析(NMDS)等方法,分析了两种生境中鲌类食物来源的差异.结果表明,两种生境中鲌类食物来源基本一致,且食物来源较为广泛,包括沿岸带饵料鱼类、敞水区饵料鱼...     

Stable resource polymorphism along the benthic littoral–pelagic axis in an invasive crayfish

Although intraspecific variability is now widely recognized as affecting evolutionary and ecological processes, our knowledge on the importance of intraspecific variability within invasive species is still limited. This is despite the fact that understanding the linkage between within‐population morphological divergences and the use of different trophic or spatial resources (i.e., resource polymorphism) can help to better predict their ecological impacts on recipient ecosystems. Here, we quantified the extent of resource polymorphism within populations of a worldwide invasive crayfish species, Procambarus clarkii, in 16 lake populations by comparing their trophic (estimated using stable isotope analyses) and morphological characteristics between individuals from the littoral and pelagic habitats. Our results first demonstrated that crayfish occured in both littoral and pelagic habitats of seven lakes and that the use of pelagic habitat was associated with increased abundance of littoral crayfish. We then found morphological (i.e., body and chelae shapes) and trophic divergence (i.e., reliance on littoral carbon) among individuals from littoral and pelagic habitats, highlighting the existence of resource polymorphism in invasive populations. There was no genetic differentiation between individuals from the two habitats, implying that this resource polymorphism was stable (i.e., high gene flow between individuals). Finally, we demonstrated that a divergent adaptive process was responsible for the morphological divergence in body and chela shapes between habitats while difference in littoral reliance neutrally evolved under genetic drift. These findings demonstrated that invasive P. clarkii can display strong within‐population phenotypic variability in recent populations, and this could lead to contrasting ecological impacts between littoral and pelagic individuals.     

Habitat Use by YOY in the Littoral Zone of an Artificially Heated Reservoir

Both temporal and spatial distributions of a young of the year (YOY) fish community were studied in the littoral zone of a French reservoir. The time survey indicated that littoral occupation highly depends on YOY time of appearance. During late spring and summer, the smallest white bream YOY (L_T < 20 mm) were restricted to complex habitats whereas those of pumpkinseed occupied all available habitats. Growing white bream progressively migrated to open water areas. At that time, pumpkinseed was the only species sampled in abundance in the littoral zone and was still collected in all habitats. The success of the introduction of pumpkinseed might be explained by both its late hatching and its opportunistic strategy in habitat selection enabling it to exploit physical spaces left vacant. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Intraspecific competition drives multiple species resource polymorphism in fish communities

It has been hypothesized that inter-specific competition will reduce species niche utilization and drive morphological evolution in character displacement. In the absence of a competitor, intra-specific competition may favor an expansion of the species niche and drive morphological evolution in character release. Despite of this theoretical framework, we sometimes find potential competitor species using the same niche range without any partitioning in niche. We used a database on test fishing in Sweden to evaluate the factors (inter- and intraspecific competition, predation, and abiotic factors) that could influence habitat choice of two competitor species. The pattern from the database shows that the occurrence of perch and roach occupying both littoral and pelagic habitats of lakes in Sweden is a general phenomenon. Furthermore, the results from the database suggest that this pattern is due to intra-specific competition rather than inter-specific competition or predation. In a field study, we estimated the morphological variation in perch and roach and found that, individuals of both species caught in the littoral zone were more deeper bodied compared to individuals caught in the pelagic zone. Pelagic perch fed more on zooplankton compared to littoral perch, independent of size, whereas the littoral perch had more macroinvertebrates and fish in their diet. Pelagic roach fed more on zooplankton compared to littoral roach, whereas littoral individuals fed more on plant material. Furthermore, we sampled littoral and pelagic fish from another lake to evaluate the generality of our first results and found the same habitat associated morphology in both perch and roach. The results show a consistent multi-species morphological separation in the littoral and pelagic habitats. This study suggests that intra-specific competition is possibly more important than inter-specific competition for the morphological pattern in the perch-roach system.     

Trophic divergence of Lake Kivu cichlid fishes along a pelagic versus littoral habitat axis

Local adaptation to the littoral and pelagic zones in two cichlid haplochromine fish species from Lake Kivu was investigated using morphometrics. Cranial variation and inferred jaw mechanics in both sexes of the two species across the two habitat types were quantified and compared. Comparisons of littoral versus pelagic populations revealed habitat‐specific differences in the shape of the feeding apparatus. Also, kinematic transmission of the anterior jaw four‐bar linkage that promotes greater jaw protrusion was higher in the pelagic zone than in the littoral zone for both species. Inferred bite force was likewise higher in pelagic zone fish. There were also sex‐specific differences in craniofacial morphology as males exhibited longer heads than females in both habitats. As has been described for other cichlids in the East African Great Lakes, local adaptation to trophic resources in the littoral and pelagic habitats characterizes these two Lake Kivu cichlids. Similar studies involving other types of the Lake Kivu fishes are recommended to test the evidence of the observed trophic patterns and their genetic basis of divergences.     

Quantifying rotifer species richness in temperate lakes

1. Biodiversity assessments of lakes depend on the ability to identify the complement of species present, although the degree of sampling required is often uncertain. We utilise long‐term data to predict rotifer species richness in three habitats in three Polish lakes using rarefaction sampling methods. 2. Richness in littoral and psammon habitats did not saturate, even with up to 130 samples. Highest richness was observed in psammon habitat (119 species) in Lake Mikolajskie, followed by littoral habitat in Lakes ?uknajno (114 species) and Kuc (110 species). Littoral habitats in Lakes ?uknajno (56%) and Kuc (51%) had the most species not shared with other habitats in the same lake. 3. Species richness (Chao2) estimates ranged between 44 for pelagic and 135 for psammon habitat in Lake Mikolajskie, to 100 for psammon and 137 for littoral habitat in Lake Kuc, and 65 for pelagic and 162 for littoral habitat in Lake ?uknajno. Whole lake estimates were 167, 205 and 171 species, respectively, for these lakes, higher than the 150 to 160 species predicted by Dumont and Segers (Hydrobiologia, 1996, 341 , 125). 4. Using standardised sampling, richness was significantly higher in littoral than either pelagic or psammon habitats. Contrasts of standardised rarefaction curves revealed that richness in Lakes Kuc and Mikolajskie was described as well by littoral‐only or psammon‐only samples, respectively, as by those randomly drawn from across all habitats in the lake. 5. Species richness estimates for Lake Mikolajskie were highest in summer, followed by autumn and spring. Interannual estimates differed by up to 427%, nearly an order of magnitude greater than maximal seasonal variation of 70%. 6. Results indicate that much higher sampling intensity is required to establish species richness than is presently carried out in most lakes. Because many species can be detected only with very intensive sampling, conservation programmes must consider sampling intensity when designing studies.     

Evidence of yellow perch, largemouth bass and pumpkinseed metapopulations in coastal embayments of Lake Ontario

Coastal embayments have been and will continue to be constructed along the northwest shoreline of Lake Ontario to restore and create warmwater fish habitat. However, very little is known about the biological connections among embayments. Using otolith microchemistry on pumpkinseed, largemouth bass and yellow perch collected from three constructed embayments in 2006?C2009, we confirm that these three species of fish each exist in a metapopulation. We find that juvenile pumpkinseed, largemouth bass and yellow perch occupy embayments different from their natal habitat after their first winter, and for at least pumpkinseed, continue to move among embayments after their second winter. We hypothesize that these fishes move among embayments after haphazardly dispersing from their overwintering habitat to the littoral zone each spring. Habitat restoration and remediation efforts in coastal Great Lakes habitats should take a system-based management approach that considers the spatial proximity of embayments, and attempts to create or preserve connected networks.     

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.     

Ecosystem response to earlier ice break‐up date: Climate‐driven changes to water temperature,lake‐habitat‐specific production,and trout habitat and resource use

Climate warming has yielded earlier ice break‐up dates in recent decades for lakes leading to water temperature increases, altered habitat, and both increases and decreases to ecosystem productivity. Within lakes, the effect of climate warming on secondary production in littoral and pelagic habitats remains unclear. The intersection of changing habitat productivity and warming water temperatures on salmonids is important for understanding how climate warming will impact mountain ecosystems. We develop and test a conceptual model that expresses how earlier ice break‐up dates influence within lake habitat production, water temperatures and the habitat utilized by, resources obtained and behavior of salmonids in a mountain lake. We measured zoobenthic and zooplankton production from the littoral and pelagic habitats, thermal conditions, and the habitat use, resource use, and fitness of Brook Trout (Salvelinus fontinalis). We show that earlier ice break‐up conditions created a "resource‐rich" littoral–benthic habitat with increases in zoobenthic production compared to the pelagic habitat which decreased in zooplankton production. Despite the increases in littoral–benthic food resources, trout did not utilize littoral habitat or zoobenthic resources due to longer durations of warm water temperatures in the littoral zone. In addition, 87% of their resources were supported by the pelagic habitat during periods with earlier ice break‐up when pelagic resources were least abundant. The decreased reliance on littoral–benthic resources during earlier ice break‐up caused reduced fitness (mean reduction of 12 g) to trout. Our data show that changes to ice break‐up drive multi‐directional results for resource production within lake habitats and increase the duration of warmer water temperatures in food‐rich littoral habitats. The increased duration of warmer littoral water temperatures reduces the use of energetically efficient habitats culminating in decreased trout fitness.     

Lipid composition in muscle and liver of sympatric coregonid fishes from Lake Baikal (<Emphasis Type="Italic">Coregonus</Emphasis> spp.) under common garden experiment

The content of total lipids, total phospholipids, and fatty acids of total lipids in muscles and liver of juvenile Lake Baikal sympatric coregonid fishes have been analyzed for the first time under a common garden experiment. Baikal omul, Coregonus migratorius Georgi, is an active migrant of the pelagic zones of the lake. Baikal (lacustrine) whitefish, C. baicalensis Dybowski, is a colonizer of the bottom habitats, which are relevant to the pelagic zones of the littoral and underwater slope. Structural lipids (phospholipids and cholesterol) dominated in total lipids of the tissues of all fish under study. Spare lipids significantly prevailed in omul muscles when compared to whitefish. The highest variability of fatty acid composition was reported in muscles of coregonid fishes. Statistically significant differences were revealed in the content of the ω3 polyunsaturated fatty acids in muscles of lacustrine whitefish and omul. Associations of lipid compositions revealed in tissues of the whitefishes under the study with their respective ecotypes have been discussed.