Density-related foraging behaviour in Closterocerus tricinctus, a parasitoid of the leaf-mining moth, Cameraria hamadryadella

• 1 We examined the foraging behaviour of the parasitoid wasp, Closterocerus tricinctus (Ashmead) (Chalcidoidea: Eulophidae), as it visited larvae of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae), in an outbreak population.
• 2 We tracked females of C.tricinctus, recording the time spent searching for mines and handling host larvae. The density of leaf-mines (host larvae) and their condition were recorded for each leaf visited. A subset of leaves visited by C.tricinctus was enclosed in fine mesh bags so that foraging success could be determined by rearing or dissection. The average density of mines and the average leaf-area mined was estimated for a random sample of leaves from each tree.
• 3 The selection of leaves upon which to forage appears to be density-dependent. C.tricinctus visits leaves with leaf-mine densities twice the average, and when switching leaves lands directly on leaf-mines 5 times more often than expected assuming random landings.
• 4 The total time spent foraging on a leaf, the average time spent handling hosts, and the total search time within leaves tend to decline on leaves with many hosts, but the observed declines are not statistically significant.
• 5 The proportion of leaf-mines visited within a leaf is strongly inversely density-dependent. 30% of visits to leaf-mines are re-visits and 29% of handling time is spent re-handling previously visited hosts. Furthermore, only 21% of visits to mines lead to successful parasitism. We suggest that self-interference and the avoidance behaviour of the host may reduce the number of visits of leaf-mines by C.tricinctus within a leaf.
• 6 The effect of the strongly inversely density-dependent foraging investment within leaves is to offset the observed density-dependent pattern of leaf visitation making the overall spatial pattern of visitation by C.tricinctus to mines of C.hamadryadella inversely density-dependent.
• 7 We suggest that the uncertainty of C.tricinctus surviving on multiply mined leaves because of density-dependent host mortality due to intraspecific competition in high-density host populations, the rarity of high-density host populations, and the rarity of multiply-mined leaves in low-density host populations combine to select against an aggregative response within leaves by C.tricinctus.

     

Lawns have vertical stratification

Abstract. Vertical stratification was examined in a closely-mown lawn, at both the beginning and end of the mowing/regrowth cycle. Two treatments were examined: Undisturbed, which had been maintained as lawn for 30 yr, and Mechanically Perturbed, on which succession was occurring, after complete removal of vegetation 9.5 months previously. There was significant vertical stratification at both stages of the mowing/regrowth cycle, i.e. even immediately after mowing to a height of 3 cm, and in both treatments. There was general consistency of the vertical role of species during the cycle and between treatments. Relative vertical positions were not closely correlated with absolute leaf heights. However, in no case was the vertical position of a pair of species significantly reversed during the cycle or between treatments. No mutually cyclic vertical relations between species were seen. It is concluded that vertical stratification occurs in even the shortest communities, it starts to develop quite early in succession, and it is dependent largely on the intrinsic properties of the species, their height and (at least in lawns) their recovery from defoliation.     

The effects of host plant phenology on the demography and population dynamics of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae)

Abstract.

• 1 We examined the effects of variation in the timing of spring leaf production and autumn leaf fall on the survival, mortality and abundance of Cameraria hamadryadella on Quercus alba and Q.macrocarpa.
• 2 We monitored and manipulated the timing of foliation on field and potted Q.alba trees and observed the abundance of C.hamadryadella on those trees. We also monitored and manipulated the timing of leaf fall on Q.alba and Q.macrocarpa trees in the field and observed its effects on survival, mortality and abundance of C.hamadryadella.
• 3 Variation in the timing of spring leaf production has no effect on C. hamudryadella abundance. However, a warm winter and spring in 1991 led to accelerated development and the imposition of a facultative third generation in one out of ten years of observation.
• 4 In 1989, leaves fell relatively early and leaf fall in the autumn accounted for more than 50% of the mortality of C.hamudryadella. in 1990 and 1991 leaves fell relatively late and leaf fall induced mortality was substantially reduced and overwinter survival was markedly increased.
• 5 The abundance of C.hamadryadella remained constant in the spring and summer of 1990 following the previous autumn's relatively early leaf fall, but increased by 10-fold in the spring of 1991 following the relatively late leaf fall of autumn 1990. The abundance of C.hamadryadella also increased 4-fold between the summer of 1991 and the spring of 1992 after another autumn of relatively late leaf fall. We attribute these increases in abundance in part to reduced mortality because of later leaf fall.
• 6 Variation in the timing of autumn leaf fall may be responsible for initiating outbreaks of C.hamadryadella.

     

Between-forest variation in vertical stratification of drosophilid populations

Abstract 1. The spatial distribution pattern of forest‐dwelling drosophilid flies was compared among species and among four forests with markedly different vertical foliage structures, including secondary and primary forests, with special reference to vertical stratification. 2. All 20 drosophilid species analysed showed vertically stratified distribution patterns, which were detected statistically in at least one forest site during the 2‐year study period. As a whole, the vertical distribution patterns were stable, with or without stratification, over the 2 years. 3. The ratio of species with stratified distribution to the total number of species changed in a similar trend with the vertical foliage complexity of forests, both showing a large gap between secondary and primary forests, even for species common to all four forests. 4. Many species, most of which were tree‐sap feeders, showed highly predictable patterns of vertical stratification across forests and years. This was associated strongly with regular preference for microhabitat, mostly for the canopy, suggesting that the unvarying nature of canopy environments in any forest is very important in producing, maintaining, and promoting the vertical stratification in organism abundance, and contributes to the ubiquity of the pattern. 5. Some species with a wider range of food resources than other species, most of which showed no clear preference for the canopy, were characterised by large between‐forest differences in vertical distribution patterns and tended to show non‐stratified patterns in secondary forests, contributing to the lower ratio of species with stratified distribution in the two secondary forests than in the primary forests. 6. The role of forest structure in organising the environment is discussed; it is suggested that vertical complexity of foliage structure affects the prevalence of vertical stratification in animal communities indirectly through the vertical heterogeneity of environmental conditions.     

Effects of the light environment on oviposition preference and survival of a leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae), on Quercus alba L.

Abstract.  1. Oviposition preference and the survival and mortality of larvae of Cameraria hamadryadella (Lepidoptera) on the foliage of Quercus alba L. (Fagaceae) in sun and shade were examined. To separate the effects of the environment of foliation from that of rearing, C. hamadryadella were allowed to oviposit on oak saplings that foliated either in the sun or shade. Subsequently, trees were placed into the sun or shade and larval survival was monitored.
2. Trees that foliated in the shade had leaves that were 30% longer than those that foliated in the sun. However, the density of leaf mines did not differ between trees that foliated in the sun or shade.
3. Larval survival was higher in the shade than in the sun, but no difference in survival due to the site of foliation was detected. Differences in larval survival between the sun and shade environments could not be attributed to differences in natural enemy mortality. However, 'death by other causes' was higher for larvae reared in the sun than in the shade. In sunny environments photochemical processes may reduce foliage quality, which results in lower survival of C. hamadryadella larvae.
4. Ovipositing C. hamadryadella do not appear to discriminate between foliage produced in the sun or shade. This lack of discrimination, coupled with the fact that ovipositing C. hamadryadella are active after dark, accounts for the absence of a difference in abundance between sun and shade.
5. The results presented here, combined with other published studies, argue that different patterns of insect herbivory and abundance in relation to the light level experienced by host-plant foliage might be expected for species that oviposit either in the night or in the day.     

Numerical modeling of vertical stratification of Lake Shira in summer

A one-dimensional numerical model and a two-dimensional numerical model of the hydrodynamic and thermal structure of Lake Shira during summer have been developed, with several original physical and numerical features. These models are well suited to simulate the formation and dynamics of vertical stratification and provide a basis for an ecological water-quality model of the lake. They allow for the quantification of the vertical mixing processes that govern not only the thermal structure but also the nutrient exchange, and more generally, the exchange of dissolved and particulate matter between different parts of the lake. The outcome of the calculations has been compared with the field data on vertical temperature and salinity distributions in Lake Shira. Lake Shira is meromictic and exhibits very stable annual stratification. The stratification is so stable because of the high salinity of the water. If the water in Lake Shira were fresh and other parameters (depth, volume, and meteorology) were the same, as now, the lake would be mixed in autumn. Using the newly developed models and using common meteorological parameters, we conclude that Lake Shira will remain stratified in autumn as long as the average salinity is higher than 3‰.     

Movement and vertical stratification of fruit-feeding butterflies in a managed West African rainforest

We analysed movement parameters and vertical stratification in fruit-feeding butterflies between a control, a thinned and a plantation site within a West African rainforest. Overall, distances moved between traps were largest in the plantation. Movement parameters were generally largest in species feeding on early successional hostplants of gap and margin habitats. In these species, distances between recaptures were significantly shorter in the thinned compared to the control forest. Conversely, forest floor species feeding on climbers and understorey shrubs showed significantly larger movement in the thinned forest. Higher strata species also flew larger distances in the thinned understorey. Including higher strata samples, they were significantly less abundant in the thinned plot, although understorey sampling alone indicated the contrary. Comparing vertical distribution patterns between thinned and control sites indicated a disruption of vertical stratification after thinning. Canopy species seem to fly in the upper strata of the more closed-canopy control forest, whereas they descend more frequently in the forest opened by the thinning management. Understorey sampling might therefore lead to biased conclusions due to differences in vertical distribution between forest plots. This study showed that thinning can affect the restricted-range forest floor butterflies as well as the more widespread canopy butterfly fauna.     

Perspectives for the biological control of Cameraria ohridella

The horse chestnut leaf-miner, Cameraria ohridella Deschka et Dimic (Lepidoptera, Gracillariidae) is a serious invasive pest of Aesculus hippocastanum in Europe. The larvae of this species feed on leaf parenchyma and can reduce the tree growth. We studied the impact of parasitoids on C. ohridella in the Czech Republic and also searched for entomopathogenic fungi associated with this pest. The results showed that the rate of parasitism varied between 5% and 15% in most cases. The most parasitized stages of C. ohridella were spinning stages and especially pupae. The most abundant parasitoid species were Minotetrastichus frontalis, Pnigalio sp. and Pediobius saulius (Hymenoptera, Eulophidae). All species are polyphagous. Using the Galleria-bait method we isolated many strains of entomopathogenic fungi. Dominant species were Paecilomyces fumosoroseus, Paecilomyces farinosus and Beauveria bassiana. The perspectives of fungal bioagents in control of C. ohridella is discussed.     

Patterns of vertical stratification in a subtropical constructed wetland in south Florida (USA)

The South Florida Water Management District (District) has built large treatment wetlands, known as Stormwater Treatment Areas (STAs), to reduce excess phosphorus loading to the Everglades. The District conducted research in a prototype treatment wetland, the Everglades Nutrient Removal Project (ENRP), to study biogeochemical processes that are important to treatment performance. Vertical profile measurements of water temperature, dissolved oxygen, pH, conductivity and transmission of photosynthetically active radiation (PAR) were made in open-water areas and sites dominated by emergent, floating or submersed vegetation over an annual cycle. Relative thermal resistance to mixing was used to infer the strength of thermal stratification. Long-term diel variation in temperature at the surface and bottom of an open-water and a vegetated site also was measured. Open-water sites were nearly isothermal and had minimal thermal stratification, while vegetated sites were all thermally stratified to some degree. The highest surface water temperatures (>35 °C) occurred in submersed vegetation where much of the light absorbed by leaves and stems was reflected as heat. Oxygen was uniformly low (<4 mg L⁻¹) in emergent and floating vegetation and attributed to shading, high biological oxygen demand and limited reaeration at the surface. Depressed oxygen levels at open-water areas were attributed to high sediment oxygen demand. The highest oxygen concentrations occurred in submersed vegetation beds. Water column pH was unstratified and near circumneutral in the open water and at emergent and floating vegetation sites, while pH was markedly stratified in submersed vegetation, where surface values at times exceeded 9.0. High surface oxygen and pH levels in submersed vegetation were consistent with intense photosynthesis. Conductivity increased at the bottom of emergent and submersed vegetation but not at open-water or floating vegetation sites. PAR transmission was strongly reduced at all sites due to shading and/or absorption by dissolved organic carbon. Light extinction coefficients were markedly higher at vegetated sites compared to the open water. Peak irradiance shifted to longer wavelengths (538–643 nm) and both short (<400 nm) and long (>700 nm) wavelengths were largely attenuated at 60 cm relative to the surface. Long-term monitoring at a vegetated site revealed periods of inverse thermal stratification and dampened diel variation in temperatures at the bottom compared to open water.     

The vertical distribution of some ciliated Protozoa in the plankton of a eutrophic pond during summer stratification

The vertical distribution of some ciliated Protozoa in the plankton of a pond in north-west England was investigated during August 1971. At this time, when the pond was stratified with an oxygen dificient hypolimnion, ciliates were counted at 10-cm depth intervals every 5 h over 25 h. The most common species (Loxodes magnus and L. striatus) were confined to the hypolimnion; there was no diurnal migration into the epilimnion. Earlier work had shown that Loxodes species require oxygen; it is therefore possible that these ciliates, which inhabited the oxygen dificient hypolimnion, migratedvertically, from time to time, to an oxygen supply at the boundary with the well-oxygenated epilimnion. To test this, Loxodes populations were confined in cellophane tubese both in the hypolimnion (at 3 m) and epilimnion (0.5 m) for 12 and 24 h (earlier trials had shown that the tubes were not markedly toxic). The ciliates died at both depths, and in a further experiment when Loxodes were confined at 3 m and 0.5 m and sampled at 5-h intervals up to 25 h it was found that they survived longer in the hypolimnion. It is suggested that ciliatees confined at 3 m died because they were unable to migrate vertically to an oxygen supply, while those at 0.5 m died because some other adverse factor was operating in the eiplimnion. Laboratory experiments showed that Loxodes died inn water in which phytoplankton photosynthesis took place and it is suggested that side effects of photosynthesis in the epilimnion (e.g. a rise in pH) caused the death of ciliates exposed at 0.5 m.     

Temporal variation in the vertical stratification of blubber fatty acids alters diet predictions for lactating Weddell seals

Fatty acid signature analysis of blubber has been used to study the foraging ecology of some marine mammals. However, species-specific information on fatty acid (FA) deposition, distribution and mobilization is required to develop further the application of FA as trophic markers within the marine environment. Blubber samples were collected from adult female Weddell seals post-parturition and end of lactation, and were divided into inner and outer half sections. We determined the degree to which there was vertical stratification in FA composition, and how this changed over the lactation period. Inner and outer layers of post-parturition blubber cores separated into two distinct groups. Sixty-two per cent of the dissimilarity between the two layers was accounted for by a higher abundance of monounsaturated fatty acids (18:1ω9c and 16:1ω7c) in the outer blubber layer, and more saturated fatty acids (16:0 and 14:0) in the inner layer. By end of lactation, the FA composition of the inner layer was different to post-parturition samples, and 20:5ω3 had the highest fractional mobilization of all FA. In contrast, the proportion of FA in the outer layer did not change, and there was more variability in the fractional mobilization of FA indicating mobilization was not uniform across the blubber layer. Dietary predictions changed considerably when highly mobilized FA were removed from analyses, and predictions were more consistent with previous dietary studies. The lack of uniformity in FA mobilization adds problems to the future use of FASA in dietary predictions, highlighting the need for more detailed information on FA mobilization.     

Analysis of vertical stratification of Syrphidae (Diptera) in an oak-hornbeam forest in northern Italy

There is growing interest in the vertical stratification of forest biodiversity, but little is known about this aspect for Syrphidae living in forests. For this study, seven Malaise traps were suspended in the canopy and seven were placed on the ground, in a lowland forest in northern Italy. Altogether, 868 specimens of Syrphidae were collected; 229 from the canopy and 639 from the ground, comprising a total of 53 species. Three species [(Ceriana conopsoides (Linnaeus), Eupeodes corollae (Fabricius), Psilota atra Fallén)] were associated significantly with the canopy layer, and seven species [(Brachypalpoides lentus (Meigen), Chalcosyrphus nemorum (Fabricius), Chrysotoxum cautum (Harris), Eumerus flavitarsis Zetterstedt, Helophilus pendulus (Linnaeus), Myathropa florea (Linnaeus) and Xylota segnis (Linnaeus)] with the ground level. The sex-ratio of the Syrphidae differed significantly between the two layers, as females were caught mainly at ground level, while males were caught preferentially in the canopy. Species with saprophagous and saproxylic larvae were caught more commonly at ground level. Species richness declined from April to November but vertical stratification was constant for most of the season. Multivariate analyses showed that two factors were correlated significantly with the community composition of Syrphidae: height of Malaise trap and estimated age of dominant trees.     

Monitoring and dispersal of the invading Gracillariidae Cameraria ohridella

The horse‐chestnut leafminer Cameraria ohridella is an invasive lepidoptera (Gracillariidae) which has spread throughout Europe over the last two decades. The species can be detected easily because of its noticeable damage to horse‐chestnut trees leaves, which allows the easy collection of large‐scale monitoring data. It is therefore an interesting model species to study invasion processes, and this review summarizes recent progresses in that regard. The date reported here result from monitoring of C. ohridella population based on pheromone traps and visual observation of damage carried out in Belgium, France, Germany, Switzerland and the UK. The patterns of spread measured using these different monitoring method, and the key factors associated with short‐distance dispersal (within cities), and large‐scale dispersal (across landscapes) are presented and discussed.     

Contrasting patterns of vertical stratification in two moth families in a Costa Rican lowland rain forest

The vertical stratification of two species-rich moth families (Lepidoptera: Arctiidae, Geometridae) was analysed in a lowland rain forest in Costa Rica. Moths were trapped with automatic 8 W ultraviolet light traps at three sites in the understorey and canopy of a primary forest at the La Selva Biological Station (10.4° N, 84.0° W). A total of 846 arctiid moths (148 species) and 946 geometrid moths (140 species) were analysed. Species richness and diversity of arctiid moths was significantly lower in the understorey than in the canopy (Fisher's alpha: 24 vs. 49) whereas geometrid moths showed an inverse pattern (Fisher's alpha: 44 vs. 27). This resulted in an overall increase in the proportion of conspicuously coloured species towards the canopy. Moth ensembles were clearly separated in multidimensional scaling ordinations, and differed significantly in their faunal composition and dominance between the strata. The available host plant data suggest that the flight height of moths was determined by larval resource availability. Examples include understorey flyers such as the geometrid genus Eois feeding on Piper, and canopy flyers such as the arctiid moth genera Aclytia, Macrocneme and Poliopastea which feed on lianas.     

Drought tolerance associated with vertical stratification of two co-occurring epiphytic bromeliads in a tropical dry forest

Vertical stratification of epiphytes generally has not been reported for dry forests. For two epiphytic Crassulacean acid metabolism bromeliads that segregate vertically, it was hypothesized that different potentials for photoprotection or shade tolerance rather than drought tolerance is responsible for the observed stratification. The light environment, capacity for photoprotection, germination response to light quality, and responses to light and drought were thus examined for Tillandsia brachycaulos and T. elongata. Vertical and light-environment distributions differed for the two species but photoprotection and photodamage did not where they occurred at similar field locations; T. brachycaulos had a higher pigment acclimation to light. Tillandsia brachycaulos had higher acid accumulation under low light as opposed to T. elongata, which responded similarly to all but the highest light treatment. Tillandsia brachycaulos maintained positive total daily net CO(2) uptake through 30 d of drought; T. elongata had a total daily net CO(2) loss after 7 d of drought. The vertical stratification was most likely the result of the sensitivity to drought of T. elongata rather than differences in photoprotection or shade tolerance between the two species. Tillandsia elongata occurs in more exposed locations, which may be advantageous for rainfall interception and dew formation.     

Forest use and vertical stratification in fruit-feeding butterflies of Sulawesi,Indonesia: impacts for conservation

Protected forest areas of Sulawesi are gradually being replaced by intensively used agroforestry systems and farmland, especially in lowland and sub-montane regions. Studies on the impact of these man-induced changes on biodiversity are of urgent conservation concern. We compared the fruit-feeding butterfly assemblage of a natural hill forest to that of a disturbed hill forest, representing a mosaic of old secondary forest and recently abandoned or active subsistence farms. Overall, species richness seemed highest in the disturbed site, but both abundance and diversity of endemic butterflies were significantly higher in the natural forest. Although the butterfly assemblage showed a clear vertical structure in the natural forest, vertical stratification was no longer pronounced at the disturbed site. Comparative studies based on diversity estimates from ground samples should consider not only the scale at which sampling is carried out and influences from nearby habitat patches in the surrounding landscape mosaic, but also possible behavioural changes in stratified species after forest modification. This study shows that higher overall species richness does not imply higher species distinctiveness, and indicates that the contribution of land-use systems to global biodiversity should be evaluated with caution, even when relatively high species richness estimates are found.     

Temporal evolution and vertical stratification of Microcystis toxic potential during a first bloom event

In order to study the setup of a Microcystis bloom and the evolution of its toxic potential, we studied the temporal and vertical variations in Microcystis aeruginosa abundance, microcystins (MC) concentrations (intracellular and extracellular), and the relative proportion of potentially microcystin-producing cells (MC-producing cells) in relation to physicochemical parameters in the recently setup Moroccan reservoir “Yaacoub Al Mansour.” The Microcystis bloom appeared relatively late in the season and was associated with a low proportion of MC-producing cells in the water surface layer, probably related to non-limiting nutrient concentrations. Interestingly, the setup of the bloom leads to a vertical gradient, showing a decrease in Microcystis cell abundance inversely coupled with an increase in the proportion of MC-producing cells. Thus, this can be the result of the growth where non-MC-producing cells remain in the lighted water layer easier than MC-producing ones. Nevertheless, parameters other than light intensity may influence the toxic potential of bloom as no vertical pattern was observed concerning microcystins cellular quotas. The high microcystins concentrations measured in the deep water layer have also proved the importance of considering the deep part of aquatic ecosystem in the management of health risks associated with cyanobacterial proliferations.     

Structure and vertical stratification of plant galler–parasitoid food webs in two tropical forests

Abstract 1. Networks of feeding interactions among insect herbivores and natural enemies such as parasitoids, describe the structure of these assemblages and may be critically linked to their dynamics and stability. The present paper describes the first quantitative study of parasitoids associated with gall‐inducing insect assemblages in the tropics, and the first investigation of vertical stratification in quantitative food web structure. 2. Galls and associated parasitoids were sampled in the understorey and canopy of Parque Natural Metropolitano in the Pacific forest, and in the understorey of San Lorenzo Protected Area in the Caribbean forest of Panama. Quantitative host–parasitoid food webs were constructed for each assemblage, including 34 gall maker species, 28 host plants, and 57 parasitoid species. 3. Species richness was higher in the understorey for parasitoids, but higher in the canopy for gall makers. There was an almost complete turnover in gall maker and parasitoid assemblage composition between strata, and the few parasitoid species shared between strata were associated with the same host species. 4. Most parasitoid species were host specific, and the few polyphagous parasitoid species were restricted to the understorey. 5. These results suggest that, in contrast to better‐studied leaf miner–parasitoid assemblages, the influence of apparent competition mediated by shared parasitoids as a structuring factor is likely to be minimal in the understorey and practically absent in the canopy, increasing the potential for coexistence of parasitoid species. 6. High parasitoid beta diversity and high host specificity, particularly in the poorly studied canopy, indicate that tropical forests may be even more species rich in hymenopteran parasitoids than previously suspected.