Colour plasticity and background matching in a threespine stickleback species pair

Examining differences in colour plasticity between closely‐related species in relation to the heterogeneity of background colours found in their respective habitats may offer important insight into how cryptic colour change evolves in natural populations. In the present study, we examined whether nonbreeding dorsal body coloration has diverged between sympatric species of stickleback along with changes in habitat‐specific background colours. The small, limnetic species primarily occupies the pelagic zone and the large, benthic species inhabits the littoral zone. We placed benthic and limnetic sticklebacks against extremes of habitat background colours and measured their degree of background matching and colour plasticity. Benthics matched the littoral background colour more closely than did the limnetics, although there was no difference between species in their resemblance to the pelagic background colour. Benthics were able to resemble both background colours by exhibiting greater directional colour plasticity in their dorsal body coloration than limnetics, which may be an adaptive response to the greater spectral heterogeneity of the littoral zone. The present study highlights how habitat‐specific spectral characteristics may shape cryptic coloration differences between stickleback species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 902–914.     

The Freshwater Zooplankton of Sri Lanka,with a Discussion of Tropical Freshwater Zooplankton Composition

Based on the study of over 500 zooplankton samples collected in Sri Lanka (Ceylon) during 1965–1974, the species composition from different habitats is analysed. The zooplankton assemblage is typically tropical with relatively few species of Cladocera and Copepoda. The Rotifera include a large number of species of the genus Brachionus. The limnetic zooplankton resembles the pond zooplankton closely in that all the eurytopic species found in the different types of habitats, including ponds, also occur in the limnetic zooplankton. The large Cladocera belonging to the genus Daphnia are very rare. In general, large zooplankters are absent. The size composition of the zooplankton has a smaller range than in temperate regions. This is due to the absence of large-sized zooplankton species. The reasons for the differences in species variety and size composition between zooplankton of temperate and tropical regions is perhaps due to a number of factors. These include the effects of high and uniform temperatures, food availability and predation by fish and invertebrates.     

Invertebrate zooplankton predator composition and diversity in tropical lentic waters

Invertebrate zooplankton predators are generally less diverse in average species numbers in tropical than in temperate lakes and reservoirs. Predatory Copepoda which comprise the majority of limnetic predators are particularly low in species numbers in the tropics. Predatory Cladocera are confined to the North Temperate zone. Chaoborus appears to be cosmopolitan. Among Rotifera, only the cosmopolitan predator Asplanchna occurs in tropical waters while the other common limnetic carnivorous genus Ploesoma is restricted to higher latitudes. Hydracarina, and insects besides Chaoborus, are generally restricted to the littoral and appear to be more diverse in the tropics. Lakes Awasa and Zwai, Ethiopia, are almost devoid of predators in the limnetic, which is invaded by a littoral chydorid Alona diaphana. Low diversity of lake types and low production of tropical zooplankton could restrict predator diversity too. Very low diversity of invertebrate predators in the limnetic and higher diversity in the littoral may characterize tropical lakes in contrast to temperate lakes, which have more invertebrate predators in the limnetic and perhaps relatively less in the littoral. Tropical zooplankton in freshwaters, appears to be a very immature community. Hence opportunistic species can readily invade the limnetic and even dominate in isolated situations as has been shown for Alona davidi, Hydracarina and some other unconventional forms.     

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.     

Community structure of fishes attracted to shallow water fish aggregation devices off South Carolina,U.S.A.

Synopsis Twenty-two fish aggregation devices were deployed in 14 m of water off South Carolina. Species composition and abundance were determined by diver visual census on eight occasions from May through November, 1985. A total of 21 families and 36 species of fishes was observed at 121 stations. Pelagic fishes dominated the fauna with a 99.3% relative abundance, and Decapterus punctatus accounted for 97.6% of the individuals. Caranx crysos, Diplectrum formosum, Decapterus punctatus, Centropristis striata and Monacanthus hispidus were the most frequent species. Total fish abundance, number of species and abundance of four of the six most common species were significantly affected by season. Hurricane activity may have caused a significant drop in pelagic fish abundance at the FADs in July. No significant correlations among species abundances were found after removal of season and FAD type effects. Spatial zonation and seasonal occurrence patterns suggest some competition among pelagic fishes. Several factors that regulate FAD faunal abundance and composition are hypothesized, including: juvenile fish availability, availability of shelter, availability of adequate food resources, interspecific and intraspecific competition, severe sea conditions, and sporadic intrusions of large predatory fishes. It is hypothesized that the abundances of benthic and pelagic FAD fishes are correlated and that there is a direct or indirect energetic link between shallow water pelagic and benthic fish assemblages near FADs.     

The Chydorid Crustacean Alona diaphana (King), Limnetic in Lakes Awasa and Zwai,Ethiopia, with a Discussion of Littoral and Limnetic Zooplankton

Alona diaphana, a common littoral tropicopolitan cladoceran, occurs in the limnetic of two Ethiopian lakes in considerable numbers and over long periods of time. Our results are presented and the composition of limnetic and littoral zooplankton, in terms of previous records of such incursions, are discussed. The high persistent densities of A. diaphana are due to the extensive littoral areas in the lakes, where it originates, the high intrinsic rates of increase of this species, the low diversity of zooplankton, reducing competition and the low level of predation.     

Phenotype-specific feeding behaviour of two arctic charr Salvelinus alpinus morphs

The feeding performance of two morphs of a trophically polymorphic fish, the arctic charr Salvelinus alpinus, feeding on zooplankton, was studied in the laboratory. Limnetic morphs, with a fusiform body, terminal mouth, short pectoral fins, and relatively long and dense gill rakers, fed at significantly higher rates on Daphnia and Cyclops than did benthic morphs. The benthic morphs were characterized by a chunky body form, subterminal mouth, long pectoral fins, and relatively short and spaced-out gill rakers. The limnetic morphs attacked the prey at significantly higher rates and were more successful in capture. Benthic morphs were more reluctant to start feeding on zooplankton and reaction time was longer than in limnetic morphs. In the absence as well as presence of zooplankton food, the position of the two morphs in the aquaria differed markedly. The results agree with dietary differences found in field-collected fish. The limnetic morphs consume zooplankton, whereas benthic morphs specialize on Lymnaea gastropods. The results suggest that benthic morphs are phenotypically inferior at using the pelagic food niche. This implies that the current role of competitive or predatory interactions in segregating the morphs may be minor.     

Sympatric and Allopatric Divergence of MHC Genes in Threespine Stickleback

Parasites can strongly affect the evolution of their hosts, but their effects on host diversification are less clear. In theory, contrasting parasite communities in different foraging habitats could generate divergent selection on hosts and promote ecological speciation. Immune systems are costly to maintain, adaptable, and an important component of individual fitness. As a result, immune system genes, such as those of the Major Histocompatability Complex (MHC), can change rapidly in response to parasite-mediated selection. In threespine stickleback (Gasterosteus aculeatus), as well as in other vertebrates, MHC genes have been linked with female mating preference, suggesting that divergent selection acting on MHC genes might influence speciation. Here, we examined genetic variation at MHC Class II loci of sticklebacks from two lakes with a limnetic and benthic species pair, and two lakes with a single species. In both lakes with species pairs, limnetics and benthics differed in their composition of MHC alleles, and limnetics had fewer MHC alleles per individual than benthics. Similar to the limnetics, the allopatric population with a pelagic phenotype had few MHC alleles per individual, suggesting a correlation between MHC genotype and foraging habitat. Using a simulation model we show that the diversity and composition of MHC alleles in a sympatric species pair depends on the amount of assortative mating and on the strength of parasite-mediated selection in adjacent foraging habitats. Our results indicate parallel divergence in the number of MHC alleles between sympatric stickleback species, possibly resulting from the contrasting parasite communities in littoral and pelagic habitats of lakes.     

Structure and dynamics of zooplankton communities in the littoral zone of some North Carolina lakes

Species composition and seasonal, dynamics of zooplankton in the littoral zone were studied in three piedmont North Carolina lakes for one year. Nygaard's compound index indicated oligotrophic conditions in Belews Lake and eutrophic conditions in Salem Lake and Lake 150. A total of 32 species of zooplankton were found in the samples. Significantly fewer species of truely littoral zooplankton were present in the oligotrophic lake regardless of season (P < 0.05). Eutrophication appears to favor increased diversity in zooplankton of the littoral zone. Habitat heterogeneity, provided by a well developed littoral zone containing aquatic macrophytes, may be the major factor contributing to the maintenance of more taxa of zooplankton in eutrophic systems.Distinct taxa of Cladocera and Copepoda are present in the littoral zone, exhibiting seasonal dynamics that are largely different from the limnetic fauna. Such dynamics may result from predation by characteristically limnetic species of cyclopoid copepods which seek prey in the littoral zone. The numerical and seasonal abundance of predators and prey are inversely related and the true littoral taxa, represented in the present study by the genera Alona, Chydorus, Pleuroxus, Sida, Simocephalus, and Eucyclops, contained no predaceous species. The seasonal dynamics of these genera are very similar even in widely separated geographical regions, indicating that the factor(s) responsible operate independently of climatic and chemical variables. Intrazooplankton predation appears to be a more plausable explanation than segregation along depth which, although consistent with data from studies of limnetic waters, cannot be related to populations of zooplankton in shallow littoral areas.     

Female mate preferences for male body size and shape promote sexual isolation in threespine sticklebacks

Female mate preferences for ecologically relevant traits may enhance natural selection, leading to rapid divergence. They may also forge a link between mate choice within species and sexual isolation between species. Here, we examine female mate preference for two ecologically important traits: body size and body shape. We measured female preferences within and between species of benthic, limnetic, and anadromous threespine sticklebacks (Gasterosteus aculeatus species complex). We found that mate preferences differed between species and between contexts (i.e., within vs. between species). Within species, anadromous females preferred males that were deep bodied for their size, benthic females preferred larger males (as measured by centroid size), and limnetic females preferred males that were more limnetic shaped. In heterospecific mating trials between benthics and limnetics, limnetic females continued to prefer males that were more limnetic like in shape when presented with benthic males. Benthic females showed no preferences for size when presented with limnetic males. These results show that females use ecologically relevant traits to select mates in all three species and that female preference has diverged between species. These results suggest that sexual selection may act in concert with natural selection on stickleback size and shape. Further, our results suggest that female preferences may track adaptation to local environments and contribute to sexual isolation between benthic and limnetic sticklebacks.     

Summer zooplankton dynamics in the limnetic and littoral zones of a humic acid lake

The limnetic and littoral zooplankton were studied during summer 1989 in Triangle Lake, a humic acid (pH 4.9) bog lake in Ohio, USA. The limnetic zooplankton showed low species richness and biomass, and dominance by the rotifers Kellicottia bostoniensis and Polyarthra vulgaris. In the littoral, species richness and biomass were markedly higher, and the crustaceans Alona guttata, Ceriodaphnia quadrangula, Chydorus sphaericus, Simocephalus serrulatus and Tropocyclops prasinus, and the rotifer Ascomorpha ecaudis were the dominants. The extreme rotifer dominance and lack of cladocerans in the limnetic zone were likely the result of Chaoborus predation. A pronounced mid-summer decline in cladoceran biomass in the littoral was likely due to predation by T. prasinus and Utricularia (bladderwort).     

Systematics and identification of larvae and juveniles of the genusSebastes

Synopsis Current information on systematics and morphology of larvae and juveniles ofSebastes is reviewed, as are intrageneric relationships among the many species of the genus. Although some subgenera may remain after a cladistic analysis of the genus, others are probably artificial groupings of species that have convergent morphologies because of similar ecology. The relationships ofSebastes to other scorpaeniform fishes as well as the relationships of scorpaeniforms to other fishes are not resolved. Of the 102 species ofSebastes worldwide, 69 species have been illustrated as preflexion larvae, 35 as postflexion larvae and 65 as pelagic juveniles. Morphological characters of each of these stages were used to compare species, group them by similar appearances and then compare these groupings with existing subgenera. Some pigment patterns of preflexion larvae were rather consistent among species within certain subgenera but quite variable among species in other subgenera. Postflexion larvae fell into 11 groups based on pigment, head spines and body shape. These groups were not closely aligned with the subgeneric assignments of the component species. On the basis of pigment patterns of pelagic juveniles, six groups were evident, but the specific composition of these groups bore little resemblance to either the larval or subgeneric groups.     

The freeliving freshwater cyclopoid copepoda (crustacea) of Malaysia and Singapore

A study of freeliving freshwater Cyclopoida of Malaysia and Singapore yielded 15 species. Some remarks are made on the morphology including a species belonging to the schmeili group of Thermocyclops. Shallow and vegetation-bearing habitats had the largest numbers of species. Most of the species are cosmopolitan. The three common species, Mesocyclops leuckarti, Thermocyclops crassus and Microcyclops varicans are eurytopic. M. leuckarti and T. crassus are the only species occurring commonly in limnetic situations in contrast to a much wider limnetic Cyclopoida species spectrum in temperate lakes. There appears to be a reduction in Cyclopoida from temperate to tropical latitudes, but the data on which this are based are at present fragmentary.     

Environmental gradients and the horizontal distribution of microcrustaceans in lakes

1. The assemblage of suspension‐feeding microcrustaceans in lakes changes along a habitat gradient from nearshore to offshore. This gradient of microcrustaceans was explored in relation to differences in macrophytes and the associated changes in water chemistry, food resources and types of predators.
2. Some microcrustacean species were littoral or limnetic specialists, while others changed their distribution along this horizontal habitat gradient on a diel or seasonal basis. Distribution patterns were similar in a lake and a pond which differed in extent of macrophyte habitat.
3. There was a large shift in the composition of sestonic food, indicating heterotrophic seston nearshore and more autotrophic seston offshore. Sit‐and‐wait predators of microcrustaceans (e.g. Enallagma spp.) dominated nearshore and cruising predators (e.g. Leptodora kindtii Focke) were more common offshore.
4. A reciprocal transplant experiment revealed that littoral specialists could survive equally well when fed littoral or limnetic seston, while limnetic specialists performed poorly when fed littoral seston. Food resources may be important in determining where some microcrustacean species live along this horizontal habitat gradient.     

Changes in Pelagic Bacteria Communities Due to Leaf Litter Addition

In many limnetic systems, the input of allochthonous organic matter, e.g., leaf litter, is a substantial source of dissolved organic carbon (DOC) for pelagic bacteria, especially in fall and winter when autochthonous DOC production is low. However, relatively little is known about community changes of pelagic lake bacteria due to leaf litter input which includes both the release of leaf leachates and microorganisms from the leaf litter into the surrounding water. Therefore, we have experimentally studied the effects of different types of leaf litter (Betula pendula, Fagus silvatica, and Pinus silvestris) on the pelagic bacterial community composition by adding leaves to different treatments of epilimnic water samples (unfiltered, 0.2 μm and 5.0 μm-pre-filtered) from humic Lake Grosse Fuchskuhle (Northeastern Germany). The addition of leaf litter led to a significant increase in DOC concentration in lake water, and each leaf litter type produced significantly different amounts of DOC (p = <0.001) as well as of specific DOC fractions (p = <0.001), except of polysaccharides. DGGE banding patterns varied over time, between types of leaf litter, and among treatments. Bacteria belonging to known bacterial phylotypes in the southwest basin of Lake Grosse Fuchskuhle were frequently found and even persisted after leaf litter additions. Upon leaf litter addition, α-proteobacteria (Azospirillum, Novosphingobium, and Sphingopyxis) as well as β-proteobacteria (Curvibacter and Polynucleobacter) were enriched. Our results indicate that supply of leaf litter DOM shifted the bacterial community in the surrounding water towards specific phylotypes including species capable of assimilating the more recalcitrant DOC pools. Statistical analyses, however, show that DGGE banding patterns are not only affected by DOC pools but also by treatment. This indicates that biological factors such as source community and grazing may be also important for shifts in bacterial community structure following leaf litter input into different lakes.     

Ontogenetic habitat shifts by Galaxias gracilis (Galaxiidae) between the littoral and limnetic zones of Lake Kanono, New Zealand

Synopsis Diel and spatial differences in distribution were determined for the larvae, juveniles, and adults of Galaxias gracilis (Galaxiidae) in a New Zealand dune lake during summer months. Larvae (mostly 10–25 mm TL) and juveniles (25–40 mm TL) inhabited shallow (0–3 m) waters of the limnetic zone and fed predominantly on two limnetic zooplankton species; Bosmina meridionalis and a calanoid copepod. At about 40 mm TL, fish moved from the limnetic to the littoral zone and expanded dietary breadth from two to over seven main prey species, including five species of littoral invertebrates. After reaching a size of about 60 mm TL, most fish moved back offshore to the deeper waters (5–15 m) of the limnetic zone during the day, moving back to the littoral zone at night to feed on invertebrates. The selection of different intra-lacustrine habitats by the various size groups of G. gracilis, and the movements between them, are interpreted as adaptive responses to the interaction between ontogenetic changes in feeding requirements and predation risk.     

Crater lake cichlids individually specialize along the benthic–limnetic axis

A common pattern of adaptive diversification in freshwater fishes is the repeated evolution of elongated open water (limnetic) species and high‐bodied shore (benthic) species from generalist ancestors. Studies on phenotype‐diet correlations have suggested that population‐wide individual specialization occurs at an early evolutionary and ecological stage of divergence and niche partitioning. This variable restricted niche use across individuals can provide the raw material for earliest stages of sympatric divergence. We investigated variation in morphology and diet as well as their correlations along the benthic‐limnetic axis in an extremely young Midas cichlid species, Amphilophus tolteca, endemic to the Nicaraguan crater lake Asososca Managua. We found that A. tolteca varied continuously in ecologically relevant traits such as body shape and lower pharyngeal jaw morphology. The correlation of these phenotypes with niche suggested that individuals are specialized along the benthic‐limnetic axis. No genetic differentiation within the crater lake was detected based on genotypes from 13 microsatellite loci. Overall, we found that individual specialization in this young crater lake species encompasses the limnetic‐ as well as the benthic macro‐habitat. Yet there is no evidence for any diversification within the species, making this a candidate system for studying what might be the early stages preceding sympatric divergence.     

Diet composition and feeding of European anchovy Engraulis encrasicolus in Kiel Bight,western Baltic Sea

Diet composition of the expanding southern species European anchovy Engraulis encrasicolus in the western Baltic Sea was investigated. Results revealed an interesting case of bentho‐pelagic coupling with potential implications for local fish species through competition for food resources.     

Some remarks on the latitudinal distribution of Cladocera on the Indian subcontinent

One hundred and thirty species of Cladocera have been recorded from the Indian subcontinent which extends from 6°N to 37°N latitude and covers an area of 4.5 million km². The equatorial region has few Daphnia species, all belonging to the sub-genus Ctenodaphnia while the more northern parts have more species of Daphni including Daphnia s. str. The limnetic Cladocera lacks the carnivorous Polyphemidae and Leptodoridae at lower latitudes (equatorial). The common limnetic species of the equatorial region are eurytopic and extend throughout the subcontinent.