Trophic ecology and persistence of invasive silver carp Hypophthalmichthys molitrix in an oligotrophic South African impoundment

The alien invasive silver carp Hypophthalmichthys molitrix established a self-sustaining feral population in an oligotrophic impoundment, Flag Boshielo Dam, in South Africa. The ability of this population to persist in a dam with low algal biomass (median annual suspended chlorophyll a = 0.08 µg l^?1), and limited access to rivers considered large enough for successful spawning, has implications for their invasive potential in other systems. Stomach content and stable isotope analysis were used to assess the trophic ecology of H. molitrix, which was then compared with indigenous Mozambique tilapia Oreochromis mossambicus, on a seasonal basis during 2011. Hypophthalmichthys molitrix are generalist filter feeders, with a diet consisting primarily of sediment, vegetative detritus, dinoflagellates and diatoms. The dominance of sediments in their stomachs suggests occasional benthic scavenging. However, H. molitrix occupied a higher trophic level (TL = 2.8) than expected, suggesting that this population subsidised their diet with an unidentified dietary constituent, characterised by enriched nitrogen values. Although the stomach contents indicated dietary overlap between H. molitrix and O. mossambicus, stable isotopes revealed fine-scale resource partitioning, despite both species occupying the same trophic level. Nonetheless, the persistence of this feral H. molitrix population in an oligotrophic impoundment highlights their phenotypic plasticity.     

Oviposition habitat selection by container‐dwelling mosquitoes: responses to cues of larval and detritus abundances in the field

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Is resource partitioning the key? The role of intra‐interspecific variation in coexistence among five small endemic fish species (Characidae) in subtropical rivers

This study focused on intra and interspecific variations in the diet of small species in the Pelotas River, upper Uruguay River ecoregion. We tested the hypothesis that five small characids coexist via resource partitioning. Samplings were performed quarterly between August 2013 and May 2014 at 14 sites in the Pelotas River basin, using gillnets of different mesh sizes, seines, cast nets and electrofishing. In this study, the contents of 1,253 stomachs were analysed from Astyanax dissensus, A. paris, A. saguazu, A. xiru and Bryconamericus patriciae. The main food resources consumed were aquatic insects, leaves, seeds and detritus. There were significant seasonal intra and interspecific variations in the use of food resources by the five species. Permutational analysis of multivariate dispersions results indicated significant differences in individual intraspecific variability between the sampled periods, with the highest values found in November and February because of an increase in allochthonous resources (leaves and seeds). Besides this, significant differences in the proportions of consumption of allochthonous and autochthonous were observed. The trophic niche breadth was significantly different among the species, such that in August, the species had narrower niche breadths. In the other periods, the highest niche breadth values were due to an increase in the consumption of allochthonous items. Diet overlap was low (< 0.4) between 62% of the species pairs. The small characid species showed pronounced changes in resource use and intra and interspecific variations indicating resource partitioning. These factors appear to be essential in species interactions and for coexistence.     

Trophic morphology features allow Astyanax endemic species coexistence in a Neotropical river system

Resource partitioning has an essential role in interspecific relations, especially in congener species, which share many morphological traits. In some places, small characids coexist through resource partitioning, which may reduce their interspecific competition. Astyanax species (e.g., Astyanax minor, Astyanax gymnodontus and Astyanax bifasciatus), for example, coexist in different water bodies from the Iguaçu River basin. These species have high phenotypic plasticity and many morphologic specializations that allow them to live in different habitats. Based on evidences that these species modified their feeding habits because of changes in resource availability in Iguaçu River, this study tested two hypotheses: (a) there are differences in head morphology, number of teeth and number of gill rakers among the species of Astyanax; and (b) there are differences in gill arch and gill raker morphology among the species of Astyanax, which may favour their trophic resource partitioning in Iguaçu River. The head morphological traits and quantitative morphological characters were summarized in a principal coordinates analysis (PCoA), and the analysis of similarities (ANOSIM) showed significant differences among species. Gill morphological measurements were analysed through analysis of covariance (ANCOVA), and it also showed significant differences in gill arch and gill raker morphology among species. Therefore, the analysis of ecomorphological traits related to trophic habits revealed some differences that may suggest a tendency of reducing competition for trophic resources in the Iguaçu River basin.     

Species diversity and decomposition in laboratory aquatic systems: the role of species interactions

1. Interest in the effects of biodiversity on ecosystem processes is increasing, stimulated by the global species decline. Different hypotheses about the biodiversity‐ecosystem functioning (BEF) relationship have been put forward and various underlying mechanisms proposed for different ecosystems. 2. We investigated BEF relationships and the role of species interactions in laboratory experiments focussing on aquatic decomposition. Species richness at three different trophic levels (leaf detritus, detritus‐colonising fungi and invertebrate detritivores) was manipulated, and its effects on leaf mass loss and fungal growth were assessed in two experiments. In the first, monocultures and mixtures of reed (Phragmites australis), alder (Alnus glutinosa) and oak (Quercus cerris) leaf disks were incubated with zero, one or eight fungal species. Leaf mixtures were also incubated with combinations of three and five fungal species. In the second experiment, reed leaf disks were incubated with all eight fungal species and offered to combinations of one, two, three, four or five macroinvertebrate detritivores with different feeding modes. 3. Results from the first experiment showed that leaf mass loss was directly related to fungal mass and varied unimodally with the number of fungi, with a maximum rate attained at intermediate diversity in oak and reed and at maximum diversity in alder (the fastest decomposing leaf). 4. Mixing litter species stimulated fungal growth but interactions between species of fungi slowed down decomposition. In contrast, mixtures of macroinvertebrate detritivores reduced fungal mass and accelerated leaf decomposition. Possible explanations of the positive relationship between detritivore diversity and decomposition are a reduction in fungal dominance and a differentiation in the use of different resource patches promoted by higher fungal diversity. 5. In conclusion, the results show a general increase in decomposition rate with increasing biodiversity that is controlled by within‐ and between‐trophic level interactions, and support the hypothesis of both bottom‐up and top‐down effects of diversity on this process.     

Resource limitation and intraspecific patterns of weight x length variation among spring detritivores

This study examined the intraspecific variability of a phenotypic trait, the body weight/body length ratio, and its adequacy to provide unbiased information about patterns of resource availability among conspecific individuals. Individual body weight and length were measured for the amphipod Gammarus minus Say (and other detritivores) in samples from freshwater springs differing in expected resource availability, and from sites in which detritus abundance had been manipulated. Mean individual weight per length was lower: (a) in populations from low-richness than from high-richness detritivore guilds; (b) in populations with size-abundance distributions similar to that of the entire guild, rather than statistically different; (c) in population samples from outside than from inside the areas of detritus addition. Small-sized individuals (< 3 mm) showed the largest variation among both populations and population samples. Similar differences were qualitatively observed for the other common detritivores co-occurring with G. minus in some springs. These observed patterns were in agreement with the variation of resource availability expected among field conditions, supporting the relevance of weight per length as a measure of food limitation.     

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.     

Gut passage times in Gammarus pulex (Crustacea,Amphipoda) and aspects of summer feeding in a stony stream

Gut passage times in Gammarus pulex (Crustacea, Amphipoda) were calculated by counting and measuring voided faecal pellets. Following feeding on elm leaves conditioned in a stream, or summer fine detritus from the same source, calculated gut passage times were usually short, about 2 h or less at 15 °C. Using harder oak and beech leaves, not previously conditioned in the stream, apparent gut passage times were much longer, 8–48 h, but since intermittent feeding occurred these figures are not comparable to the others.In connection with summer feeding of the animal in the stream, the microbiological status of the fine detritus there was studied. It contained fungal hyphae, which were mostly empty, viable fungal spores of terrestrial derivation, and bacteria, all largely carried on fragments of vascular plant tissues. In experiments, such fungal spores (of Aureobasidium pullulans and of Mucor sp.), incorporated into fine detritus, were not digested by the animal. The bacterium Bacillus cereus was present in the fine detritus and when the latter was consumed it survived passage through the gut of the animal. Using B. cereus as a proportion marker it was concluded that other unicellular bacteria in the fine detritus were not digested either. In other experiments, not immediately related to the summer season, nutrient extraction by the animal from hyphae of the fungus Nectria lugdunensis was examined; it probably occurs through sub-microscopic pores connecting the cells. This mode of extraction may apply when the animal consumes vascular plant tissues.     

CYST–THECA RELATIONSHIP,LIFE CYCLE,AND EFFECTS OF TEMPERATURE AND SALINITY ON THE CYST MORPHOLOGY OF GONYAULAX BALTICA SP. NOV. (DINOPHYCEAE) FROM THE BALTIC SEA AREA1

A new species of Gonyaulax, here named Gonyaulax baltica sp. nov., has been isolated from sediment samples from the southeastern Baltic. Culture strains were established from individually isolated cysts, and cyst formation was induced in a nitrogen‐depleted medium. Although G. baltica cysts are similar to some forms attributed to Spiniferites bulloideus and the motile stage of G. baltica has affinities with G. spinifera, the combination of features of cyst and motile stage of G. baltica is unique. The culture strains were able to grow at salinity levels from 5 to 55 psu and formed cysts from 10 to 50 psu. Cultures at each salinity level were grown at 12, 16, and 20° C. Temperature‐ and salinity‐controlled morphological variability was found in the resting cysts. Central body size varied with temperature and salinity, and process length varied with salinity. Cysts that formed at extreme salinity levels displayed lower average process length than cysts formed at intermediate salinity levels, and central body length and width were lowest at higher temperature and lower salinity. Models for the relationship between central body size and temperature/salinity and process length and salinity have been developed and may be used to determine relative paleosalinity and paleotemperature levels. Our results on salinity‐dependent process length confirm earlier reports on short‐spined cysts of this species found in low salinity environments, and the model makes it possible to attempt to quantify past salinity levels.     

Seasonal variations in food resource partitioning among four sympatric gudgeon species in the upper Yangtze River

Knowledge of food resource partitioning among sympatric fish species is crucial for understanding the potential mechanisms of species coexistence. Gudgeons (Teleostei: Cyprinidae: Gobioninae) often dominate fish assemblages in the upper Yangtze River. However, little research has been conducted on their trophic interactions. In this paper, seasonal diet and feeding strategy variations of four sympatric gudgeon species, Coreius guichenoti, Coreius heterodon, Rhinogobio ventralis, and Rhinogobio cylindricus, were investigated by analysis of intestinal tract contents, aiming to explore whether food resource partitioning occurred among them. Fish specimens were collected during spring (April–May) and autumn (August–October) in 2010 in Hejiang, a free‐flowing stretch of the upper Yangtze River. Coreius guichenoti, C. heterodon, and R. cylindricus showed omnivorous feeding habits, while R. ventralis exhibited an obligate carnivore feeding habit. Diet overlap among the four studied species was high, especially in spring. However, changes in feeding strategies were observed in autumn. Specifically, C. guichenoti and R. cylindricus expanded their dietary niche breadth and consumed detritus, Sinopotamidae or Hydropsychidae as important complementary food resources. In contrast, C. heterodon and R. ventralis reduced their dietary niche breadth and became more specialized on mussels (Limnoperna lacustris). These results confirmed that sympatric fish species can coexist with high diet overlap, and food resource partitioning among these species may also fluctuate with the seasons.     

The role of the freshwater shrimp <Emphasis Type="Italic">Atyaephyra desmarestii</Emphasis> in leaf litter breakdown in streams

In aquatic ecosystems, microorganisms and invertebrates provide critical links between plant detritus and higher trophic levels. Atyaephyra desmarestii is an omnivorous decapod that inhabits freshwaters and exhibits high tolerance to temperature oscillations and high ability to colonize new habitats. Although A. desmarestii is able to ingest a variety of foods, few studies have been conducted to elucidate the role of this freshwater shrimp on detritus breakdown in streams. In this study, A. desmarestii was allowed to feed on conditioned or unconditioned alder and eucalyptus leaves in microcosms with or without access to its fecal pellets. At the end of the experiment, total body length of the animals was measured, and the remaining leaves and fecal pellets were used for dry mass quantification and assessment of bacterial and fungal diversity by denaturing gradient gel electrophoresis (DGGE). Cluster analyses of DGGE fingerprints indicated that the major differences in microbial communities on leaves were between leaf types, while on fecal pellets were between conditioned and unconditioned leaves. However, the consumption rate by the shrimp did not differ between leaf types, and was significantly higher on leaves conditioned by microorganisms and in treatments without access to feces. In treatments without access to feces, the production of feces and fine particulate organic matter was also significantly higher for conditioned leaves. Overall, our results support the feeding plasticity of A. desmarestii and its potential role in plant litter breakdown in streams. This might have implications for maintaining stream ecosystem functioning, particularly if more vulnerable shredders decline.     

Colour polymorphism and genetic variation in Idotea baltica populations from the Adriatic Sea and Baltic Sea

Phenotypic and genetic variation was studied in two of the four European subspecies of the marine isopod Idotea baltica; the Mediterranean I. b. basteri and the Baltic I. b. baltica. Spatial and temporal patterns of colour polymorphism were analysed in northern Adriatic and western Baltic Sea populations. Pronounced differences in phenotype composition were observed between populations of both subspecies as seen in the distribution of various colour variants bilineata, lineata, flavafusca and several combined forms). Compared with Adriatic samples, western Baltic Sea populations show higher phenotypic diversity. To obtain an estimate of the degree of genetic divergence between the subspecies, 12 gene-enzyme systems were investigated electrophoretically. The results obtained indicate a relatively high level of genetic variation; I. b. basteri from the nothern Adriatic tends to be more polymorphic and more heterozygous than I. b. baltica from the western Baltic. Both subspecies share identical electrophoretic mobilities of the homologous enzyme proteins examined; however, in allelic composition they exhibit significant differences at approximately half the number of loci scored. The genetic distance (Nei's D) measured at the subspecific level was 0.04. Amounts and geographical patterns of variation, observed both in colour phenotype and electrophoretic variation, are considered.     

Assortative pairing in Gammarus insensibilis (Amphipoda) infected by a trematode parasite

We have investigated the influence of Microphallus papillorobustus (Trematoda) on the reproductive biology and mating patterns of its intermediate host Gammarus insensibilis (Amphipoda). Infected Gammarus species show altered behaviour which renders them more susceptible to predation by Charadriiform birds, the parasite's definitive hosts. In a natural population of G. insensibilis, mean parasite intensity was higher for unpaired individuals than for paired individuals. Fecundity was reduced in infected amphipods. Size-assortative pairing was significant, although infected males were found with smaller females compared to uninfected males of the same size. There was also a positive assortative pairing by parasitic prevalence. Vertical segregation between infected and uninfected individuals, male-male competition for access to uninfected females, and female choice may explain assortative mating for prevalence. This study provides the first empirical evidence that parasites can have a direct effect on patterns of mating in gammarids.     

Swimming with the giant: coexistence patterns of a new redfin minnow Pseudobarbus skeltoni from a global biodiversity hot spot

Ecological niche theory predicts that coexistence is facilitated by resource partitioning mechanisms that are influenced by abiotic and biotic interactions. Alternative hypotheses suggest that under certain conditions, species may become phenotypically similar and functionally equivalent, which invokes the possibility of other mechanisms, such as habitat filtering processes. To test these hypotheses, we examined the coexistence of the giant redfin Pseudobarbus skeltoni, a newly described freshwater fish, together with its congener Pseudobabus burchelli and an anabantid Sandelia capensis by assessing their scenopoetic and bionomic patterns. We found high habitat and isotope niche overlaps between the two redfins, rendering niche partitioning a less plausible sole mechanism that drives their coexistence. By comparison, environment–trait relationships revealed differences in species–environment relationships, making habitat filtering and functional equivalence less likely alternatives. Based on P. skeltoni's high habitat niche overlap with other species, and its large isotope niche width, we inferred the likelihood of differential resource utilization at trophic level as an alternative mechanism that distinguished it from its congener. In comparison, its congener P. burchelli appeared to have a relatively small trophic niche, suggesting that its trophic niche was more conserved despite being the most abundant species. By contrast, S. capensis was distinguished by occupying a higher trophic position and by having a trophic niche that had a low probability of overlapping onto those of redfins. Therefore, trophic niche partitioning appeared to influence the coexistence between S. capensis and redfins. This study suggests that coexistence of these fishes appears to be promoted by their differences in niche adaptation mechanisms that are probably shaped by historic evolutionary and ecological processes.     

Behavioural and ultrastructural studies on the sedentary platyctenean ctenophore Vallicula multiformis

Feeding habit and diet composition of two colonizing mullids, Upeneus asymmetricus and U. moluccensis, and two indigenous goatfish Mullus barbatus and M. surmuletus (Mullidae), in the eastern Mediterranean were investigated. The most important taxonomic group in the diet of all four species was the Decapoda which were identified to the specific level, enabling precise assessment of trophic partitioning. Leptochela pugnax, a pasiphaeid of Red Sea origin, was by far the most important prey of all mullids studied, which resulted in considerable trophic overlap between all of the species. Trophic separation does not play an important role in resource partitioning of the eastern Mediterranean mullids. Consequently, coexistence between colonizing and indigenous mullids is presumably achieved by spatial segregation.     

Trophic interrelationships between the exotic Nile tilapia, <Emphasis Type="Italic">Oreochromis niloticus</Emphasis> and indigenous tilapiine cichlids in a subtropical African river system (Limpopo River,South Africa)

The stable isotope ratio and seasonal changes in diet of two indigenous (Oreochromis mossambicus, Tilapia rendalli) and one exotic (Oreochromis niloticus) tilapiine cichlids in the subtropical Limpopo River, South Africa were investigated to determine patterns of resource partitioning. Stomach contents of O. niloticus and O. mossambicus indicated high dietary overlap across size class, habitat and season, with both species primarily feeding on vegetative detritus. However, stable isotope analysis revealed that the two Oreochromis species had different stable isotope ratios derived from different food sources. The relatively δ¹³C-depleted O. niloticus indicates a phytoplankton-based diet, while the δ¹³C-enriched O. mossambicus indicates a macrophagous diet dominated by vegetative detritus and periphyton. The high similarity in stomach contents and the interspecific differences in isotopic composition reveal fine-scale patterns of food resource partitioning that could be achieved through selective feeding. Tilapia rendalli was largely macrophagous and fed mainly on aquatic macrophytes and had a low dietary overlap with both O. niloticus and O. mossambicus. In the Limpopo River, detritus and algae are probably the most abundant food resources and the causal factors responsible for the observed patterns of resource partitioning among the tilapiines are usually difficult to ascertain. Fish may be able to perceive food resources in terms of the dynamics that determine their availability. Detailed studies of variation in food resource availability and fish habitat use within the system are needed to evaluate this hypothesis.     

Oribatid mites reveal that competition for resources and trophic structure combine to regulate the assembly of diverse soil animal communities

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Pollution effects on the phytoplankton-zooplankton relationships in an inshore environment

The quantitative and qualitative relationships between the phytoplankton and zooplankton populations were examined for one year in an inshore environment (Elefsis Bay) of the Aegean Sea. It was found that quantitative relationships among these populations were influenced by the trophic level of the environment and their stocks were significantly correlated only in non-polluted conditions. The quantitative relationship between the principal grazers in the zooplankton population (Acartia clausi, Oithona nana) and the phytoplankton dominant species Exuviaella baltica, upon which these copepods were believed to graze preferentially, was tested and the results were discussed.     

Alternative trait combinations and secondary resource partitioning in sexually selected color polymorphism

Resource partitioning within a species, trophic polymorphism is hypothesized to evolve by disruptive selection when intraspecific competition for certain resources is severe. However, in this study, we reported the secondary partitioning of oviposition resources without resource competition in the damselfly Ischnura senegalensis. In this species, females show color polymorphism that has been evolved as counteradaptation against sexual conflict. One of the female morphs is a blue‐green (andromorph, male‐like morph), whereas the other morph is brown (gynomorph). These female morphs showed alternative preferences for oviposition resources (plant tissues); andromorphs used fresh (greenish) plant tissues, whereas gynomorphs used decaying (brownish) plants tissues, suggesting that they chose oviposition resources on which they are more cryptic. In addition, the two‐color morphs had different egg morphologies. Andromorphs have smaller and more elongated eggs, which seemed to adapt to hard substrates compared with those of gynomorphs. The resource partitioning in this species is achieved by morphological and behavioral differences between the color morphs that allow them to effectively exploit different resources. Resource partitioning in this system may be a by‐product of phenotypic integration with body color that has been sexually selected, suggesting an overlooked mechanism of the evolution of resource partitioning. Finally, we discuss the evolutionary and ecological consequences of such resource partitioning.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.