Timing is everything? Phenological synchrony and population variability in leaf‐chewing herbivores of Quercus

Abstract.  1. Specialization on ephemeral resources (e.g. new leaves) should produce large annual variation in herbivore population size when the timing of availability of those resources is unpredictable. Despite considerable evidence for impacts of synchrony with budburst on survival of larval Lepidoptera, previous studies of adult Geometridae and Noctuidae found no correlations between insect phenology and population variability.
2. We surveyed larval Lepidoptera feeding on Quercus alba and Q. velutina in Missouri from 1993 to 2003 and examined population variability, measured as the coefficient of variation of population density (CV), in a subset of abundant species. We compared CV values among species whose larvae feed only in spring, early summer, mid-summer, late summer, or all season. We predicted that univoltine species whose larvae eclose and complete development in spring during leaf expansion would have higher variability than species feeding later in the season, having multiple generations, or having longer development times.
3. As predicted and consistent with hypotheses, spring-feeding species had CV values 32% higher than species feeding in summer months. Coefficients of variation were also 34% higher in leaf-rolling and mining guilds compared with free-feeders, suggesting that mobile species may compensate for asynchrony with budburst by dispersing to higher quality plants or plant parts. Multivoltine species, however, did not differ from univoltine species in population variability.
4. Our results suggest that asynchrony with plant phenology and factors that might exacerbate it, such as climate change, will have the largest impacts on the dynamics of spring-feeding Lepidoptera, particularly species with limited mobility.     

The effect of dispersal ability in winter and summer stoneflies on their genetic differentiation

Abstract.  1. Plecopteran species disperse less than most other aquatic insects. Within stoneflies, members of different families vary in the degree of wing morphology and season of adult emergence.
2. Dispersal limitations were tested to determine if there were increased differences among the nearby, Chagrin and Grand River populations in north-eastern Ohio, by comparing genetic variation within the 16s rRNA region of mitochondrial DNA in two stoneflies. Allocapnia recta emerges in winter and often has rudimentary wings, and Leuctra tenuis emerges in summer with fully developed wings.
3. There was significant genetic variability between the samples of A. recta from two adjacent rivers (FST = 0.20), but not between samples of L. tenuis (FST = 0.07).
4. Distinct clades in A. recta were found to occur within the minimum spanning tree specific to the Chagrin River, which contributed to a significant difference in gene diversity between the two rivers. Haplotypes in L. tenuis appeared randomly distributed between the two rivers; however, nucleotide diversity was significantly less in samples from the Grand River.
5. Shared haplotypes of both species illustrate the inter-connectedness of the Chagrin and Grand River populations, and the lower genetic variability of L. tenuis between the two rivers is indicative of its greater dispersal capability.     

Altitudinal and environmental variation in lifespan in the Copper butterfly Lycaena tityrus

1.  Variation in longevity within and between natural populations is widespread, and understanding the relative importance of environmental and genetic factors as well as their interactions in mediating such variation is crucial in longevity research.
2.  In this study lifespan of adult copper butterflies was examined in relation to altitude, temperature (20 and 27 °C), sex and adult feeding.
3.  As expected, longevity increased with decreasing temperature, and sucrose-fed butterflies had longer lifespans compared to water-fed and finally non-fed individuals. The impact of feeding, especially of having access to water or not, was larger at the higher compared to the lower temperature.
4.  Regarding altitudinal patterns, increased lifespan in high-altitude populations was largely restricted to beneficial feeding conditions, while under carbohydrate deprivation low-altitude animals lived longer, suggesting that low-altitude butterflies do better under food stress.
5.  Differences in longevity between sexes were small at 20 °C, while females lived substantially longer than males at the higher temperature. Consequently, females may be less susceptible to high temperature stress than males. Further, males suffered more from food stress than females, suggesting that females are generally more stress resistant than males.
6.  Using a full factorial design, this study demonstrates that variation in longevity is caused by several factors, and additionally by substantial interactive effects. Consequently, patterns of variation in longevity are complex, and one needs to be cautious when neglecting this source of variation, by focussing on individual factors only.     

Risk-sensitive decisions during nesting may increase maternal provisioning capacity in the subsocial shield bug Parastrachia japonensis

Abstract 1. Females of the monophagous shield bug Parastrachia japonensis Scott provision their nymph-containing nests with high-quality drupes of the single host tree, Schoepfia jasminodora , a resource that is poor and unstable, under an array of variable environmental constraints.
2. Although the number of drupes provisioned is correlated positively with enhanced nymphal development and survival, there is great variability in the number of drupes that females provide.
3. The ability to adjust behaviour and/or physiology, i.e. make risk-sensitive decisions, should be adaptive for organisms such as P. japonensis that are confronted with extreme variability in food availability and in the conditions under which they must forage. To assess whether females of this species use risk-sensitivity to improve their provisioning success, data on the parameters of provisioning activity obtained during a long-term field study were analysed. The relationships between variation in three female-controlled factors during nesting (nest site, active stage during the provisioning season, duration of provisioning activity) and the conditions of three environmental factors (drupe availability, intraspecific competition, weather) were examined. The relationships between all of these factors and provisioning capacity of females were investigated.
5. Nest site, intraspecific competition, and timing of provisioning activity only affected provisioning capacity when drupe availability was extremely good or bad or weather was particularly bad.
6. The findings suggest that females use risk-sensitive decisions to increase provisioning capacity under extreme conditions of low and high drupe availability and inclement weather.     

Drought stress differentially affects leaf-mining species

Abstract.  1. The impact of climate change on phytophages is difficult to predict, due in part to variation between species in their responses to factors such as drought stress. Here, the hypothesis that several species within the leaf-mining feeding guild will respond in a consistent way to changes in rainfall patterns is tested, using a manipulative field experiment.
2. Summer drought, enhanced summer rainfall, and control treatments were imposed on a calcareous grassland community, and the responses of five leaf-mining species were assessed.
3. One leaf-mining species was more abundant under enhanced rainfall, one was more abundant under drought, and the other three species showed no consistent response to the rainfall treatments. Higher parasitism levels under drought may partly explain the response of one species ( Stephensia brunnichella ) to the treatments.
4. These results show that generalisations relating to drought stress impacts cannot be drawn at the feeding guild level for leaf-mining insects.     

Hydrological connectivity drives patterns of macroinvertebrate biodiversity in floodplain rivers of the Australian wet /dry tropics

1. Floodplain rivers in Australia's wet/dry tropics are regarded as being among the most ecologically intact and bio-diverse lotic ecosystems in the world, yet there have been relatively few community-based studies of their aquatic fauna.
2. To investigate relationships between hydrological connectivity and biodiversity in the region, macroinvertebrates were collected from sites within two contrasting floodplain rivers, the 'tropical' Gregory River and 'dryland' Flinders River systems, during the dry season and analysed at various spatial scales. A subset of sites was re-sampled in the following dry season to explore temporal variation. The fauna consisted of 124 morphotaxa, dominated by gatherers and the Insecta.
3. As predicted, hydrological connectivity (the lotic or lentic status of waterbodies) had a major influence on macroinvertebrate assemblage composition and diversity, both in space and time. Assemblages from waterbodies with similar connection histories were most alike, and beta-diversity between assemblages was greatest between lotic and lentic waterbodies, tending to increase with increasing spatial separation.
4. At smaller spatial scales, a number of within-waterbody, habitat and water quality characteristics were important for explaining variation (61%) in the taxonomic organization of assemblages, and characteristics associated with primary productivity and habitat diversity were important for explaining variation (45%) in the functional organization of assemblages. However, much of the small-scale environmental variation across the study region appeared to be related to broad-scale variation in hydrological connectivity, which had both direct and indirect effects on macroinvertebrate assemblages.
5. Conservation of the biodiversity in Australia's wet/dry tropics may depend on conserving the natural variation in hydrological connectivity and the unregulated flow of floodplain rivers.     

Can life-history traits predict the response of forb populations to changes in climate variability?

1.  Climate change will cause changes in average temperature and precipitation as well as increased fluctuations around the mean, yet few studies have considered the impacts of altered climate variability on plant populations. We tested whether life-history traits (expected life span, generation time and seed size) can predict plant responses to increased environmental variability across similar plant species sharing the same habitat.
2.  We combined long-term demographic data on 10 prairie forb species with stochastic demography techniques to estimate the effects of potential changes in matrix element means and variances on the long-term stochastic population growth rate.
3.  For all 10 species, recruitment had higher contribution and elasticity values than survival, meaning that climate change is more likely to influence population growth through effects on recruitment than on survival for these relatively short-lived forbs. Species with longer generation times had lower elasticities to increases in matrix element variability.
4.   Synthesis. Our analysis of a unique, long-term data set suggests that longer-lived plant species will be less vulnerable to the effects of future increases in climate variability. While this relationship was previously reported for diverse taxa from many locations, our results show that it also applies within a guild of short-lived species from a single community. The generality of the pattern demonstrates the potential for using life-history traits to make predictions about which species may be the most vulnerable to climate change.     

Role of dissolved organic carbon in the attenuation of photosynthetically active and ultraviolet radiation in Adirondack lakes

1. We surveyed eighty-five lakes located in the Adirondack Mountain Region of New York State, U.S.A., to characterize the attenuation of photosynthetically active (PAR) and ultraviolet radiation (UVR) in relation to dissolved organic carbon (DOC) concentrations and pH. Attenuation of PAR was quantified in situ . Attenuation was also inferred by measuring the light absorption of filtered lake water samples at wavelengths (300, 340 and 440 nm) representing UV-B, UV-A and PAR.
2. Substantial variation in transparency was observed among lakes in this region. Attenuation depths ( z _1%) for PAR ranged from 0.5 to greater than 20 m, while inferred values for UV-B and UV-A ranged from a few centimetres to > 5 m. Median values of UV-A penetration (0.75 m) and UV-B penetration (0.45 m) corresponded to 11% (UV-A) and 6% (UV-B) of lake maximum depth.
3. Much of the variation in PAR and UVR attenuation was explained by differences in lake DOC. Univariate power models based solely on DOC accounted for 85% (PAR), 90% (UV-A) and 91% (UV-B) of the variation in absorption.
4. Attenuation and absorption coefficients were generally lower for recently acidified lakes compared to acidic and circumneutral lakes which have not undergone recent acidification. However, differences among these three groups of lakes were not statistically significant. Our results suggest that the effects of acidification on the optical properties of a regional population of lakes, even in an area experiencing widespread acidification, are relatively subtle in comparison with other factors contributing to inter-lake variability.
5. The presence of near-shore wetlands is probably a key factor influencing regional variability in DOC and light climate among Adirondack lakes. Temporal variability in climatic factors influencing wetland DOC production and export may mask more subtle influences on lake DOC associated with anthropogenic acidification.     

Control of reproduction and a survival cost to mating in female Bicyclus anynana butterflies

Abstract.  1. Given that sexual conflict is all pervasive, investigating potential costs to mating and the control of female reproduction are important issues.
2. Here, female reproductive output and longevity are investigated in relation to mating status (virgin vs. once-mated females) and host-plant availability in the tropical butterfly Bicyclus anynana .
3. Both factors significantly affected realised fecundity in a manner demonstrating that female reproduction is not entirely under control of the female herself, but is rather subject to additive contributions of the female, her male partner, and intrinsic physiological processes. As evident from the deposition of significant egg numbers by virgin, host-deprived females, B. anynana is effectively unable to completely inhibit oogenesis.
4. Mated females suffered a reduction in adult life span, which cannot be explained as a side-effect of variation in egg size, lifetime, or early fecundity.
5. Such detrimental effects of mating per se are indicative of the cooperation–conflict balance between sexes being shifted towards conflict in B. anynana .     

Divergent patterns of photosynthetic phosphorus-use efficiency versus nitrogen-use efficiency of tree leaves along nutrient-availability gradients

1.  Nitrogen (N) and phosphorus (P) are essential nutrients for photosynthetic carbon assimilation and most frequently limit primary productivity in terrestrial ecosystems. Efficient use of those nutrients is important for plants growing in nutrient-poor environments.
2.  We investigated the pattern of photosynthetic phosphorus-use efficiency (PPUE) in comparison with photosynthetic nitrogen-use efficiency (PNUE) along gradients of P and N availability across biomes with 340 tree and shrub species. We used both total soil N and P concentration and foliar N/P ratios for indicating nutrient-availability gradients.
3.  Photosynthetic phosphorus-use efficiency increased with greater leaf mass per area (LMA) toward decreasing P availability. By contrast, PNUE decreased with greater LMA towards decreasing N and P availability.
4.  The increase in PPUE with decreasing P availability was caused by the net effects of a relatively greater reduction in foliar P concentration and a relatively constant photosynthetic carbon assimilation rate. The decrease in PNUE with decreasing N availability was caused by the effects of a reduction in photosynthetic carbon assimilation rate with greater LMA.
5. Synthesis . Our results suggest that higher PPUE may be an effective leaf-level adaptation to P-poor soils, especially in tropical tree species. Future research should focus on the difference between PPUE and PNUE in relation to leaf economics, physiology and strategy.     

Physico-chemistry and aquatic insects of a glacier-fed and a spring-fed alpine stream

1. Physico-chemical conditions and benthic macroinvertebrates were studied in two adjacent alpine streams in the Tyrolean Alps, Austria, for 2 years, and aquatic insect emergence was recorded for 1 year.
2. In the spring-fed system, maximum discharge and increased concentrations of suspended solids, nitrate and particulate phosphorus occurred during snowmelt in June. In the glacier-fed stream, high discharge and strong diel fluctuations in flow and concentrations of suspended solids created a harsh and unstable environment during summer. Glacial ablation, variation in groundwater inflow, and water inputs from tributaries draining calcareous rocks caused water chemistry to vary both seasonally and longitudinally in glacier-fed Rotmoosache.
3. A total of 126 aquatic or semi-aquatic invertebrate taxa were collected, 94 of which were found in the glacier-fed stream and 120 in the spring-fed stream. Chironomid abundance was 2–8 times and taxa richness 2–3 times lower in the glacier-fed stream than in the spring-fed stream, as was the number of chironomid taxa (72 versus 93 total).
4. These results broadly support the conceptual model by Milner & Petts (1994) concerning glacier-fed stream systems. However, single samples and seasonal means showed relatively high invertebrate abundance and richness, especially during winter, indicating a considerable degree of spatial and temporal variability.
5. We suggest that the seasonal shifts from harsh environmental conditions in summer to less severe conditions in autumn and a rather constant environment in winter are an important factor affecting larval development, life-history patterns and the maintenance of relatively high levels of diversity and productivity in glacier-fed streams.     

Inter-hemispheric comparison of fire history: The Colorado Front Range,U.S.A., and the Northern Patagonian Andes,Argentina

Fire history was compared between the Colorado Front Range (U.S.A.) and northern Patagonia (Argentina) by dating fire-scars on 525 Pinus ponderosa and 418 Austrocedrus chilensis, respectively, and determining fire weather on the basis of instrumental and tree-ring proxy records of climatic variation. Years of above average moisture availability preceding fire years, rather than drought alone, is conducive to years of widespread fire in the Colorado Front Range and the northern Patagonian study areas. Above-average precipitation promotes fire by enhancing the growth of herbaceous plants which increases the quantity of fine fuels during the fire season a few years later. The short-term variability in moisture availability that is conducive to widespread burning is strongly related to El Niño Southern Oscillation (ENSO) activity. The warm (El Niño) phase of ENSO is associated with greater moisture availability during the spring in both regions which leads to peaks in fire occurrence several years after El Niño events. The warmer and drier springs associated with la Niña events exacerbate the drying of fuels so that fire years commonly coincide with La Niña events. In both regions, there was a dramatic decline in fire occurrence after the early 1900s due to a decline in intentionally set fires by Native Americans and European settlers, fuel reduction by livestock grazing, and increasingly effective organized fire suppression activities after the 1920s. In both regions there was a marked increase in fire frequency during the mid-and late-19th centuries which coincides with increased ignitions by Native Americans and/or European settlers. However, year-to-year variability in ring widths of Pinus ponderosa and Austrocedrus chilensis also increased from relatively low values in the late 1700s and early 1800s to peaks in the 1850s and 1860s. This implies frequent alternation of years of above and below average moisture availability during the mid-19th century when the frequencies of major fire years rise. The high correlation of tree-growth variability betweem the two regions implies a strong inter-hemispheric variation in climatic variability at a centennial time scale which closely parallels a variety of proxy records of ENSO activity. Based on the relationship of fire and ENSO events documented in the current study, this long-term trend in ENSO activity probably contributed to the mid- and late-19th century increase in fire spread in both regions. These similar trends in fire occurrence have contributed to similar patterns of forest structures, forest health, and current hazard of catastrophic wildfire in the Colorado Front Range and northern Patagonia.     

Interdemic variation of cannibalism in a wolf spider (Pardosa monticola) inhabiting different habitat types

Abstract.  1. Cannibalism was investigated in the wolf spider Pardosa monticola (Clerck) using spiders collected from four populations with varying densities, inhabiting two different coastal dune habitat types. Sampled individuals were paired randomly and tested immediately for their cannibalism propensity.
2. The occurrence of cannibalism was found to be influenced by the size (cephalothorax width) of both the smaller and the larger spider of a pair. Larger size differences enhanced cannibalism.
3. Cannibalism rates were not significantly different in spiders from high-density compared with low-density populations. Cannibalism rates showed, however, large variability between habitat types, with higher rates in spiders from dune grasslands than from dune slacks. This is suggested to result from differences in prey availability throughout the growing season between both habitat types.
4. Different size classes of spiders did not use different microhabitats, indicating that microhabitat segregation as a cannibalism-avoidance behaviour is absent in this species.     

Genetic population structure and dispersal in Atlantic Island caddisflies

SUMMARY 1. Population genetic structure of Wormaldia tagananana, a caddis with a narrow geographic range and endemic to the Canary Islands, was investigated by studying allozyme variation at 11 putative loci in five of the eight extant populations on Tenerife and La Gomera. Genetic variability, population structure and gene flow were compared with those reported previously in more widespread Trichoptera, particularly Canarian populations of the non-endemic limnephilid Mesophylax aspersus, to examine the hypothesis that the Wormaldia , with its restricted range, would exhibit relatively little genetic variability and gene flow.
2. Despite it being a narrow-range island endemic, genetic variability in populations of W. tagananana is broadly similar to values reported for more widespread caddis.
3. Significant genetic population structure was observed in W. tagananana (overall F _ST = 0.387), greater than that seen in M. aspersus and amongst the highest reported for lotic caddis to date. Several site- and island-specific alleles were reported, providing further evidence for the relative isolation of individual Wormaldia populations.
4. Significant deviations from Hardy–Weinberg equilibrium were found in four of five populations (overall F _IS = 0.675). This could result from within-locality population substructuring, or offspring within a reach being the product of a limited number of matings.
5. This genetic evidence supports the hypothesis that the restricted range of W. tagananana is, at least in part, because of limited dispersal ability.     

Differential defoliation of Eucalyptus grandis arises from indiscriminant oviposition and differential larval survival

1 The influence of six open-pollinated families (OPFs) of Eucalyptus grandis on both the growth and development of larvae and the oviposition preference of a paropsine chrysomelid ( Paropsis atomaria ) was investigated. The OPFs had previously been identified as differing in their susceptibility to defoliation by P. atomaria in forestry progeny trials.
2 Oviposition preference for resistant and susceptible foliage was tested using binary choice tests. These tests did not demonstrate any significant preference for either resistant or susceptible open-pollinated material indicating that adult host preference for susceptible trees was not a likely cause of differential defoliation.
3 Quantification and analysis of growth and development parameters for all larval stages of P. atomaria showed that feeding on genetic material identified as resistant resulted in a significant reduction of relative growth rate of first instar larvae and an alteration to normal feeding behaviour. There was also a trend towards increased larval mortality on resistant E. grandis .
4 We argue that although the magnitude of these effects was minor, interactions with additional biotic and abiotic sources of mortality in the field have the potential, when magnified over successive generations, to result in significant variation in defoliation of host genotypes in the field.     

Resistance of Enterobacter sakazakii (Cronobacter spp.) to environmental stresses

Aim:  To gain a better understanding of the survival and persistence of Enterobacter sakazakii in severe environments.
Methods and Results:  We evaluated the resistance of Ent. sakazakii to various environmental stresses, including heating, drying, water activity ( a _w), and pH. The resistance of Ent. sakazakii to heat varies widely among strains. Most tested strains of Ent. sakazakii exhibited unusual resistance to dry stress, which depends on drying media. Growth of most strains occurred within 24 h at 37°C when the initial a _w of the medium was adjusted to 0·94 with sucrose or sodium chloride. The minimum pH for growth within 24 h at 37°C was 3·9 or 4·1 for most strains tested. Additionally, there did not appear to be any relationship between resistance to stresses and biofilm-forming ability in Ent. sakazakii planktonic cells.
Conclusions:  These results indicate that Ent. sakazakii is much more resistant than other Enterobacteriaceae to environmental stresses. Moreover, it is likely that Ent. sakazakii has cross-resistance to dry and thermal stresses.
Significance and Impact of the Study:  The findings of this study will contribute to an improved understanding of the survival and behaviour of Ent. sakazakii , which will lead to improved strategies for preventing outbreaks of Ent. sakazakii infection.     

Over-wintering stage polymorphism of a bruchine beetle: geographical variation in optimal diapause strategy

Abstract.  1. Most temperate insects undergo diapause at a specific developmental stage to overcome severe winters. The bruchine beetle Bruchidius dorsalis in a warmer region in Japan, however, has diverse over-wintering stages – non-diapausing larvae, diapausing larvae, and diapausing adults, whereas in a cooler region, it over-winters only as the final instar larva or adult in diapause.
2. The geographical pattern of over-wintering stages in 12 populations was investigated over a wide range along the mainland of Japan. It revealed that a distinct geographical pattern of over-wintering stages exists in relation to temperature conditions. In regions with warmer climates, the proportion of non-diapausing larvae increased and B. dorsalis had a more complex over-wintering stage structure.
3. Life cycles were also compared between two areas of Japan by field experiments. In the cooler area, the first generation over-wintered in the diapausing larval or adult stage. Conversely, in the warmer area, diapause was induced later and some of the first-generation adults produced second-generation offspring before over-wintering.
4. Based on the geographical cline of climates and the differences in cold hardiness among stages, we can demonstrate that the over-wintering stage variation among and within populations results from an adaptive timing of diapause induction in each region, because the late larval or adult diapauses protect pupae or eggs – which unlike other stages are not cold hardy – from being produced late in the season.     

Factors influencing size inequality in peatland black spruce and tamarack: evidence from post-drainage release growth

1 We used tree ring analysis to determine stem radius and thus examine size variation over time in two even-aged (approximately 40-year-old) mixed populations of black spruce and tamarack established on peatlands in a boreal forest. We also followed the response of one of these populations to improved edaphic conditions over 8 years following drainage.
2 Populations of trees in undrained areas showed a decline in size variability over time until age 20–25 years, after which size heterogeneity was relatively stable.
3 For trees in undrained areas there was a relationship between age and size for the first 20–25 years, but this relationship then broke down due to a period where relative growth rate and size were inversely related.
4 For the population of trees in the drained area, smaller trees (i.e. those that had been growing more slowly prior to drainage) showed significantly greater drainage-induced release growth, while larger trees (those growing faster prior to drainage) showed an initial reduction in growth following drainage and, overall, less release growth.
5 The response of tamarack to drainage was more dramatic than for black spruce.
6 Despite extensive variation in tree size, drainage dramatically reduced variability in growth rate among trees of each species, such that size variability in the populations declined.
7 We postulate that the heterogeneity of microsites with respect to edaphic conditions, perhaps associated with the hummock to hollow microtopographic gradient, has a major influence on growth variation, and hence size inequality, in peatland populations of black spruce and tamarack.     

Effects of piscivore-mediated habitat use on growth, diet and zooplankton consumption of roach: an individual-based modelling approach

SUMMARY 1. We used an individual based modelling approach for roach to (i) simulate observed diel habitat shifts between the pelagic and littoral zone of a mesotrophic lake; (ii) analyse the relevance of these habitat shifts for the diet, activity costs and growth of roach; and (iii) quantify the effects of a hypothetical piscivore-mediated (presence of pikeperch) confinement of roach to the littoral zone on roach diet, activity costs and growth.
2. The model suggests that in the presence of pikeperch, roach shifts from zooplankton as the primary diet to increased consumption of less nutritious food items such as macrophytes, filamentous algae and detritus.
3. The growth of roach between May and October was predicted to be significantly higher in the absence of pikeperch, although the net activity costs were about 60% higher compared with the scenario where pikeperch were present.
4. These modelling results provide quantitative information for interpreting diel horizontal migrations of roach as a result from a trade-off between food availability and predation risk in different habitats of a lake.
5. Altering the habitat selection mode of planktivorous roach by piscivore stocking has the potential to reduce zooplankton consumption by fish substantially, and could therefore be used as a biomanipulation technique complementing the reduction of zooplanktivorous fish.