Presence of predatory wasps and stinkbugs alters foraging behavior of cryptic and non-cryptic caterpillars on plantain (Plantago lanceolata)

We examined the foraging patterns of two species of caterpillar (Junonia coenia: Nymphalidae and Spilosoma congrua: Arctiidae) that contrast in feeding specialization and crypticity on plantain (Plantago lanceolata) in the absence and presence of two different insect predators [stinkbugs, Podisus maculiventris (Pentatomidae) and wasps, Polistes fuscatus (Vespidae)]. Junonia larvae were quite apparent to human observers, feeding on upper leaf surfaces during daylight, whereas Spilosoma larvae were relatively cryptic, often hiding under leaves and in soil crevices during daylight. In the presence of either predator species, the non-cryptic Junonia caterpillars more quickly left the plant on which they were initially placed and were less apparent than Junonia larvae not exposed to predators. The presence of predators had no detectable influence on where the caterpillars occurred on the plants (new, intermediate-aged or mature leaves, or reproductive stalks). Surprisingly, the predators influenced the behavior of the inherently cryptic Spilosoma: the apparency of these larvae at night increased when wasps had access to the plots during the day. Survivorship of the non-cryptic Junonia was less than 12% when stinkbugs were present compared to 60% in their absence. Although the presence of wasps resulted in a lower relative growth rate for the non-cryptic Junonia larvae, the indirect effect of predators on reduction in survivorship due to alterations in prey growth rate through behavioral changes was less than 3%. After taking into account the decline in caterpillars per plot through predation, we found that both the amount of leaves eaten and the proportion of plants eaten were altered on plots with predators present, which suggests that the caterpillars' increased consumption countered increased maintenance costs due to the presence of predators. Overall, our results indicate that hostplant size, level of predation and type of predator can influence the degree to which these caterpillars react to the presence of insect predators. In contrast, degree of inherent feeding specialization and cryptic behavior seemed to have little effect on the expression of reactive behaviors of these caterpillars to predators.     

Mast-seeding in the cycad genus Encephalartos: a test of the predator satiation hypothesis

Mast-seeding behaviour was monitored in 18 populations of eight species of the African cycad genus Encephalartos between 1988 and 1991. The coefficient of variation (V) in annual cone production for each population ranged between 88 and 200, indicating large fluctuations in reproductive effort between years. Data were collected to determine whether mast-seeding reduced levels of predispersal seed predation by satiating seed predators in mast years and whether it resulted in a reproductive advantage over plants which reproduced more frequently. Masting intensity was greatest in those populations in which individual plants suffered the highest levels of predispersal seed predation in years when only a few plants produced seeds. The principal seed predators were two congeneric weevil species, Antliarhinus zamiae and A. signatus, which develop exclusively on cycad seeds. The lowest intensity of mast-seeding was recorded for cycad populations with low levels of seed predation and in which A. zamiae and A. signatus occurred only in low numbers or were entirely absent. Larger seed crops appeared to result in lower levels of seed predation by A. zamiae and A. signatus in four populations of E. altensteinii, and differences in seed crop size accounted for 48–66% of variation in levels of seed predation in populations of five cycad species. In one population of E. altensteinii, lower levels of seed predation in plants reproducing periodically resulted in a reproductive advantage over plants reproducing more frequently. These results are consistent with the predator satiation hypothesis. However, in most cycad populations, numbers of seed predators did not appear to decrease significantly after a period of 2–8 years between reproductive episodes and, in two of three populations examined, periodic reproduction did not increase the number of seeds surviving to dispersal over a 4-year period. These results are interpreted to mean that periodic reproduction has not evolved in response to selection imposed by seed predators, but that selection may favour those plants which experience lower levels of seed predation by coning in synchrony with the majority of plants in the population.     

Effects of habitat fragmentation on the three-way interaction among ants,aphids and larvae of the giant purple emperor, <Emphasis Type="Italic">Sasakia charonda</Emphasis> (Hewitson), a near-threatened butterfly

Survival rates of both early and middle instar larvae of the nymphalid butterfly, Sasakia charonda, were estimated to be lowest on test trees planted in a meadow (site A), intermediate in a small, narrow secondary deciduous broadleaf forest (small patch, site B) and highest in a large secondary deciduous broadleaf forest (large forest, site C). The larval mortality rates due to predation by tree-climbing predators from the ground (tree climbing predator) such as ants and the larvae of carabids were estimated to be greater at sites A and B than those at site C. The number of predatory ants climbing test trees was significantly greater at sites A and B than at site C, and the ants harvested honeydew from aphids living on tree leaves at those two sites. Aphid densities were significantly higher on trees at sites A and B than at site C, and aphid densities and numbers of predatory ants were significantly and positively correlated at sites A and B. In an experiment controlling aphid density per branch on test trees, the numbers of ants and the mortality rates of S. charonda larvae were greater on branches with high aphid densities than on those with low aphid densities at both sites A and B. These results suggest that the aphid density per host tree was higher in the meadow and the small patch than in the large forest; at both sites these higher aphid densities attracted higher numbers of predatory ants to test trees, and as a result, mortality rates of S. charonda larvae were increased.     

Predators mediate host plant resistance to a phytophagous pest in cabbage with glossy leaf wax

This study examined the role of generalist predators in producing higher mortality ofPlutella xylostella L. (Plutellidae) larvae on glossy vs. normal-wax cabbage,Brassica oleracea var.capitata L. To test this, survival and feeding ofP. xylostella were measured on individually caged glossy and normal-wax plants with and without each of three generalist predators,Chrysoperla carnea (Stephens) (Chrysopidae),Orius insidiosus (Say) (Anthocoridae), andHippodamia convergens Guerin-Meneville (Coccinellidae). In the greenhouse, predators always significantly reduced survival ofP. xylostella larvae on glossy plants, but never on normal-wax plants. In the field, predators significantly reducedP. xylostella survival on glossy plants, but onlyC. carnea was effective on normal-wax plants. In similar experiments with excised leaves,O. insidiosus andC. carnea were more effective predators on the glossy leaves, whileH. convergens was equally effective on both kinds of leaves. Patterns for feeding were similar, but significance levels differed from those forP. xylostella survival. The greater effectiveness of predators on glossy plants is apparently due to the reported improved mobility of these animals on glossy leaf surfaces. The data also suggest that reduced mining byP. xylostella exposes the larvae to more predation on glossy plants and contributes some to the resistance. Regardless of the mechanism, resistance toP. xylostella on glossyB. oleracea appears to depend on the action of generalist predators for its full expression. This dependence on predation must be considered in the development and deployment of glossy insect-resistantB. oleracea.     

Predator Presence Moves <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> Larvae to Distraction

Displacement of herbivorous insects by the presence of predators on whole plants has rarely been studied. By semi-continuous observations of an externally feeding insect herbivore and a predator, we show how the mere presence of the predator, Geocoris lubra Kirkaldy (Hemiptera: Geocoridae), on a plant can have a strong influence on the movement and behaviors of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) larvae. The presence of predators, as opposed to mortality by predators, influenced the proportion of larvae feeding, resting and implementing avoidance activities. In addition, the proportion of time individual larvae allocated to feeding, resting and dropping off plants was affected when predators were present with and without contact between the two. Predators do more than just reduce numbers of herbivores; they influence feeding, displacement and subsequently the distribution of plant damage.     

Prey utilization by Chaoborus punctipennis Say in a small,eutrophic reservoir

Prey selection by 2nd-, 3rd-, and 4th-instar larvae of Chaoborus punctipennis was evaluated by comparing larval crop contents to the species structure of the zooplankton community in a small, eutrophic lake. The rotifers, Brachionus caudatus and Trichocerca similis, were positively selected by and comprised most of the diet of 2nd- and 3rd-instar larvae. Rotifers comprised 67% of all prey items identified in the crops of even 4th-instar larvae. Whereas 2nd-and 3rd-instar larvae selected the two rotifers with equalfrequency, 4th-instar larvae selected Trichocerca significantly more frequently than they did Brachionus. Although copepods comprised only 8% of all prey identified in 3rd-instar crops, they comprised 33% of the crop contents of 4th-instar larvae. Both 3rd and 4th instars exhibited a positive electivity for Cyclops vernalis significantly more frequently than for Diaptomus pallidus. We conclude that the larvae of Chaoborus punctipennis can, and do, select between Trichocerca and Brachionus and between Cyclops and Diaptomus.The paper by Brooks and Dodson (1965) triggered a flurry of studies on aquatic predators and their ability to discriminate between available prey species. Although the original work dealt with vertebrate predators, it has become increasingly apparent that invertebrate predators may also exhibit considerable selectivity in choosing between prey categories. The larval stages of the various species of Chaoborus are ubiquitous lentic predators, especially in mesotrophic or eutrophic systems. Although there have been a number of recent publications which deal with the food habits and preferences of several species of Chaoborus, only a few studies have included analyses of larval crop contents and none of these have included Chaoborus punctipennis. This species is of particular interest because it is one of the smaller species in the genus, and perhaps for this reason, because it is frequently present in lakes which support large populations of planktivorous fish (von Ende 1979). The purpose of our research was to characterize the food habits and preferences of Chaoborus punctipennis in a small, eutrophic reservoir which supports a rather depauperate number of species of zooplankters.     

Variation in selection pressures on the goldenrod gall fly and the competitive interactions of its natural enemies

Summary Larvae of the tephritid fly Eurosta solidaginis induce ball-shaped galls on the stem of tall goldenrod, Solidago altissima. Survival probability depends on gall size; in small galls the larva is vulnerable to parasitoid oviposition, whereas larvae in large galls are more frequently eaten by avian predators. Fly populations from 20 natural old fields in central Pennsylvania were monitored in 1983 and 1984 to examine the distribution of the selection intensity imposed by natural enemies, the parasitoids Eurytoma gigantea and E. obtusiventris, the inquiline Mordellistena unicolor, and the predatory birds Dendrocopus pubescens and Parus atricapillus. Mordellistena and E. obtusiventris are able to attack galls of all diameters while E. gigantea and the predatory birds preferentially assaulted small and large diameter galls, respectively. Eurosta in intermediate sized galls had the highest survivorship, hence selection had a stabilizing component. However, parasitoid attack was more frequent than bird attack, and the two did not exactly balance, thus there was also a directional component. The mean directional selection intensity on gall size was 0.21 standard deviations of the mean, indicating that larger gall size was favored. Interactions among the insect members of the Eurosta natural enemy guild are complex and frequent.     

Populations of Predaceous Natural Enemies Developing on Insect-Resistant and Susceptible Tomato in North Carolina

Naturally occurring populations of immature and adultGeocoris punctipes,adultColeomegilla maculataand immature coccinellids were monitored on field-grown tomato lines susceptible and resistant toManduca sextaandHelicoverpa zea. Helicoverpa zeaandHeliothis virescenseggs and small larvae that serve as prey for these predators also were monitored. MoreH. zeaandH. virescenseggs and small larvae were found on resistant than on susceptible plant lines. However, similar populations of largeH. zeaandH. virescenslarvae were found on resistant and susceptible plants. The number of adultGeocoris punctipes,adultColeomegilla maculataand immature coccinellids on resistant plants was always as high or higher than the number on susceptible plants. The data demonstrate no incompatibility of host-plant resistance with biological control provided by these predaceous insects, but indicate that the number ofG. punctipesand coccinellids required to provide effective biological control may develop too late in the season to be of practical value. Large populations of stilt bugs (Jalysus wickhami,Hemiptera: Berytidae) and spiders were observed to occur earlier in the growing season than eitherG. punctipesor coccinellids and may be a significant source of mortality forH. zeaeggs and small larvae.     

Larval parasitoids and pathogens of the groundnut leaf miner,Aproaerema modicella (Lep.: Gelechiidae), in India

Natural enemies of the groundnut leaf miner,Aproaerema modicella (Deventer), were studied at the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT) located near Hyderabad in peninsular India. Hymenopterous parasitoids attacking leaf miner larvae were the most important group of natural enemies. Nine primary and eight secondary parasitoids emerged from host larvae, and killed up to 50% of the leaf miner larvae sampled. The trophic relationships between primary and secondary parasitoids are incompletely understood. The influence of pathogens of this species is reported for the first time. These pathogens killed up to 30% of the leaf miner larvae. The combined effects of all mortality agents killed up to 95% of the leaf miner larvae per sample period. However, use of insecticides in sprayed plots reduced the efficacy of parasitoids. The impact of predators on larval populations was not studied and may explain underestimates of leaf miner mortality rates.      

Size structure and substitutability in an odonate intraguild predation system

Interactions between different size classes of predator species have the potential to influence survival of prey species in intraguild predation (IGP) systems, but few studies test for these effects. Using a substitutive design in a field setting, I measured the effects of two size classes of IG predators (large and small larvae of the dragonfly Anax junius) on the mortality of IG prey (larvae of the dragonfly Pachydiplax longipennis). I also examined whether combinations of large A. junius and P. longipennis and small A. junius and P. longipennis had substitutable effects on shared prey (larvae of the damselfly Ischnura verticalis). The presence of both size classes of A. junius, when alone and in combination with P. longipennis, significantly increased mortality of I. verticalis. In the presence of P. longipennis, large and small A. junius had similar effects on the mortality of I. verticalis, and effects of size-structured assemblages of A. junius were similar to the effects of each size class alone at the same density. The effects of the two size classes of A. junius on P. longipennis differed, and P. longipennis mortality was lower when exposed to size structured assemblages of A. junius than when exposed to only large A. junius at the same density. Results were similar to those in a laboratory study, although the effect of P. longipennis on I. verticalis was much lower in the field setting. These results demonstrate that interactions between different size classes of IG predators promote the survival of IG prey and highlight the importance of within-species size structure as a characteristic that may promote the coexistence of predators in IGP systems.     

Searching clusters of community composition along multiple spatial scales: a case study on aquatic invertebrate communities in bamboo stumps in West Timor

We examined spatial heterogeneity at multiple scales in composition of the aquatic invertebrate communities in bamboo stumps in a mountainous area of West Timor. We partitioned the study area (ca. 15,000 m²) at five levels of patchiness: two sites, four sub-sites, eight super-clumps, 14 clumps, and 84 stumps. Similarity of community composition between stumps varied more than expected from independent occurrence of each taxon in comparisons within any levels of patches. Negative association was frequently detected among taxa. These results indicate heterogeneity in community composition at a stump level. At higher levels, similarity among stumps within each site was greater than expected from null models which assumed no spatial heterogeneity, and similarities among super-clumps, sub-sites and between sites in a whole area were lower than expected from the null models. The observed patterns in similarities among subsets of the community and distribution of each taxon were mostly consistent with the models which assumed site-level heterogeneity. Therefore, we conclude that the community in this area was spatially heterogeneous at stump and site levels. The relationship between mean intra- and inter-specific crowding suggested that the site level habitat heterogeneity might reduce the chance of encounters between two predators, the larvae of the Toxorhynchites mosquito and the Brachyceran fly.     

The importance of intraguild interactions to the combined effect of a parasitoid and a predator on aphid population suppression

Intraguild predation (IGP) occurs when consumers competing for a resource also engage in predatory interactions. A common type of IGP involves aphid predators and parasitoids: since parasitoid offspring develop within aphid hosts, they are particularly vulnerable to predation by aphid predators such as coccinellid beetles. Other intraguild interactions that include non-lethal behavioral effects, such as interference with foraging and avoidance of IGP, may also hamper parasitoid activity and reduce their effectiveness as biological control agents. In this study, we quantified mortality in and behavioral effects on Aphidius colemani Viereck (Hymenoptera: Aphidiidae) by its IG-predator Coccinella undecimpunctata L. (Coleoptera: Coccinellidae), and compared the impact of two release ratios of these natural enemies on aphid populations. Parasitoids did not leave the plant onto which they were first introduced, regardless of the presence of predators, even when alternative prey was offered on predator-free plants nearby. In 2-hour experiments, predator larvae interfered with wasp activity, and the level of aphid parasitism was lower in the presence of predators than in their absence. In these experiments, the parasitoids contributed more to aphid mortality than the predators and aphid suppression was higher when a parasitoid acted alone than in combination with a predator larva. These results were confirmed in a 5-day experiment, but only at one parasitoid:predator release ratio (4:3) not another (2:3). The over-all impact on aphid population growth was non-the-less stronger when both enemies acted together than when only one of them was present. Results indicate that for given release ratios and time scale, the negative lethal and non-lethal effects of the predator on parasitoid performance did not fully cancelled the direct impact of the predator on the aphid population.     

广州小斑螟发生与环境因子的关系

广州小斑螟是红树林的一种灾害性的食叶害虫.通过室内饲养和野外观察,对广州小斑螟的发生和环境因素的关系进行了详细的研究.结果表明,随着龄级的增加,取食量增大;在不同样地不同滩位的虫口密度差异性规律不同;在单株白骨壤的不同方位虫口密度差异显著,正南方向虫口密度最高,正西、正北虫口密度最低;在单株白骨壤的中上部明显高于下部;广州小斑螟大龄幼虫较耐水淹,水淹6h的死亡率为0;不同地区温度的差异可导致广州小斑螟的发育进度的不同.     

Assessment of three techniques for measuring natural enemy inflicted mortality of leafroller larvae in commercial orchards

Measuring natural enemy (NE) inflicted mortality presents a serious challenge in managed habitats, such as apple orchards, where the crop-plant is frequently treated with a wide range of insecticides. Toxic residues keep natural populations of pests at too low levels to sample and prevent the use of techniques in which laboratory reared insects are released directly on foliage and used to measure NE activity. Three methods to implant obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) larvae were tested: (1) larvae placed on potted apple trees, (2) larvae placed on excised leaves inserted in florists’ waterpicks and (3) larvae placed on branches placed into water filled Polyvinyl chloride (PVC) pipes. Use of potted trees allowed accurate estimation of NE inflicted mortality; however, this method was very labor intensive. The waterpick method required less labor, but it underestimated larval mortality caused by NEs. The use of PVC pipes to hold infested branches combined the benefits of the other two methods tested. This technique was simple to use and provided accurate estimates of NE inflicted mortality.     

Impacts of biological control and invasive species on a non-target native Hawaiian insect

The potential for classical biological control to cause unintended harm to native species was evaluated in the case of the endemic Hawaiian koa bug, Coleotichus blackburniae White (Hemiptera: Scutelleridae), and parasitoids introduced to Hawaii for control of an agricultural pest, the southern green stink bug, Nezara viridula (L.) (Hemiptera: Pentatomidae). Parasitism of C. blackburniae eggs, nymphs and adults by biocontrol agents was quantified across a wide range of habitats and compared to other sources of mortality. Egg mortality due to the biocontrol agent Trissolcus basalis Wollaston (Hymenoptera: Scelionidae) was low (maximum 26%) and confined to elevations below 500 m on a single host plant. Predation, mainly by alien spiders and ants, was the greatest source of egg mortality (maximum 87%). Parasitism of adult C. blackburniae by the biocontrol agent Trichopoda pilipes (F.) (Diptera: Tachinidae) was near zero at 21 of 24 sites surveyed. Three sites with high bug density had higher levels of T. pilipes parasitism, reaching maxima of 70% among adult female bugs, 100% among males and 50% among fifth instars. Male-biased parasitism indicated that T. pilipes is adapted to using male aggregation pheromone for finding C. blackburniae hosts. The relative impacts of biocontrol agents and other sources of mortality were compared using life tables. Invasive species, particularly generalist egg predators, had the greatest impacts on C. blackburniae populations. Effects of intentionally introduced parasitoids were relatively minor, although the tachinid T. pilipes showed potential for large impacts at individual sites. In retrospect, non-target attacks by biological control agents on C. blackburniae were predictable, but the environmental range and magnitude of impacts would have been difficult to foresee.     

Balancing risks? Responses and non-responses of mayfly larvae to fish and stonefly predators

Summary In a series of laboratory experiments we examined the hypothesis that larvae of stream mayflies would respond to the presence of two different types of predators in such a way as to minimize their risk of being consumed by each. Positioning of larvae (whether they frequent the top, sides, or bottom of stones) of Baetis tricaudatus and Ephemerella subvaria was altered by the presence of predaceous stoneflies (Agnetina capitata) with a larger proportion of the population occurring on the upper surfaces, where the probability of encountering the predator was lowest. The presence of a benthivorous fish (Cottus bairdi) had no significant effects on positioning of the mayfly larvae. Lack of fish effects may reflect an inability of the mayflies to detect or respond to sculpins, or alternately may indicate that sculpins do not normally present a important predation risk for these mayflies. Failure of mayfly prey to account for fish predators when responding to the presence of stoneflies appcars to explain facilitation previously observed between stoneflies and sculpins.     

Non-random distribution patterns of leaf miners on oak trees

Summary Leaf-mining Stilbosis quadricustatella larvae are distributed non-randomly within leaves of their host plants, sand live oak (Quercus geminata) and water oak (Q. nigra), in north Florida. Fewer mines are found together on the same side of the mid-vein than separated, on opposite sides of the mid-vein. Larvae do not normally cross the mid-vein but create small blotch-like mines along subsidiary veins. Investigations of the usual mortality factors acting on these leaf-miner populations, including competition, parasitism, and predation, revealed no significant differences in these factors between mines separated by the mid-vein and those on the same side of the leaf. However, early leaf abscission, which kills the larvae present in the leaf, occurs significantly more frequently in cases where larvae are clustered on one leaf side. The reasons for this differential leaf abscission are not yet clear.     

Influence of Predator Presence and Prey Density on Behavior and Growth of Damselfly Larvae (Ischnura elegans) (Odonata: Zygoptera)

Foraging behavior is often determined by the conflicting benefits of energy gain and the risk of mortality from predation or other causes. Theory predicts that animals should have lower activity levels when either the risk of predation or the availability of resources in the environment is high. We investigated the adjustment of the behavior of I. elegans larvae to predator presence (Anax imperator) and prey density (Daphnia sp.) and their interaction in a completely crossed factorial experiment in the lab and the effect of behavior on growth. The foraging activity of the I. elegans larvae was significantly reduced in the presence of a free-swimming predator but not a caged predator. Abdominal movements were significantly reduced at a low prey density. Growth was significantly reduced by the presence of a free swimming predator and low prey densities. These results provide evidence that these damselfly larvae adjust their behavior to the presence of predators to increase their survival at the expense of reduced growth and development.     

Larval interactions in aphidophagous predators: effectiveness of wax cover as defence shield of Scymnus larvae against predation from syrphids

Small-sized predators in the aphidophagous guild of Aphis gossypii Glover colonies on hibiscus trees in Japan exploit aphids at low prey abundance. Scymnus (Pullus) posticalis Sicard beetles were the first predatory species to attack aphids in the spring, and their larvae co-occurred with larvae of Eupeodes freguens (Matsumura) syrphids in aphid-infested leaves of hibiscus for 3 weeks in absence of large-sized coccinellid predators. Larval interaction between Scymnus and syrphid predators was examined in relation to effectiveness of wax cover of Scymnus against predation from syrphids. Waxless first instar larvae were not protected but wax-covered larvae of second, third and fourth instars were protected from predation by syrphid larvae. The protection was lower in the second instar which has a thin wax cover and significantly higher in the third and fourth instars having a thick wax cover. In addition, larvae from which the wax was removed were significantly more vulnerable to predation. Vulnerability of Scymnus larvae to predation from syrphids was directly related to the thickness of wax cover. Results suggest that the wax cover of Scymnus larvae act as an effective defence mechanism against predation from syrphid larvae.     

Predatory behaviors of Neoseiulus californicus and Galendromus helveolus (Acari: Phytoseiidae) attacking Oligonychus perseae (Acari: Tetranychidae)

Predatory behaviors of Neoseiulus californicus (McGregor) and Galendromus helveolus (Chant) attacking Oligonychus perseae Tuttle, Baker and Abbatiello on avocado leaves were videotaped and analyzed. Behaviors were recorded for “fresh” predators that were used ≤ 48 hr post receipt from a commercial insectary and “cold stored” predators that were maintained at 12°C for ≈ 14 days. Fresh and cold stored G. helveolus were observed to attack O. perseae only after invading webbed nests. Conversely, fresh and cold stored N. californicus employed three different modes of predatory attack: (1) intercepting and attacking migrant O. perseae outside of web nests; (2) attacking prey through nest webbing; or (3) invading and attacking O. perseae inside nests. Predatory efficacy of both N. californicus and G. helveolus was reduced following cold storage, as both species engaged in certain predatory behaviors less frequently in comparison to predators that were not stored at low temperatures. Our observed results for N. californicus and G. helveolus attacking O. perseae are interpreted in relation to the chaetotaxy hypothesis, which proposes that phytoseiid invasion efficiency and propensity of webbed nests is facilitated by dorsal setal lengths. This revised version was published online in July 2006 with corrections to the Cover Date.