The species-area relationship in the hoverfly (Diptera, Syrphidae) communities of forest fragments in southern France

The effect of forest fragmentation was studied in hoverfly communities of 54 isolated forests (0.14–171 ha) in south west France. The positive relationship between species richness and wood patch area was investigated by testing the three hypotheses usually put forward to explain it: 1) the sampling effect hypothesis, 2) the patch heterogeneity hypothesis, 3) the hypothesis of equilibrium between distance from other patch (colonisation) and surface area of the patch (extinction). The syrphid species were divided into 3 ecological groups, based on larval biology as summarized in the "Syrph the Net" database: non forest species, facultative forest species and forest species. A total of 3317 adults belonging to 100 species, were captured in the 86 Malaise traps. Eight species were non forest (N=16), 65 facultative forest (N=2803) and 27 forest species (N=498).
Comparison of the slopes of the species-area curves for species richness and species density per forest patch showed a strong sampling effect in the species-area relationship. Wood patch heterogeneity increased with wood patch area and positively influenced hoverflies richness. Less isolated wood patches presented high richness of forest species and low richness of non forest species. Only forest species richness seemed to respond to the equilibrium between surface area and isolation. Depending on which hypothesis explained best the species-area relationship, management recommendations to mitigate fragmentation effects were formulated at various spatial scales and for different stakeholders.     

Zoogeographical analysis of the hoverfly fauna (Diptera, Syrphidae) of the northwestern Caucasus

The hoverfly fauna of the northwestern Caucasus comprises 257 species of 72 genera. They possess 32 types of ranges that could be united into 11 groups of ranges. Widespread species from the basis of the hoverfly fauna in the region: species with narrower ranges, those distributed in more than one zoogeographical region, and those distributed in more than one dominion, constitute 12.9%, 63.1%, and 24% of the fauna, respectively. The hoverfly fauna of the northwestern Caucasus is composed of nemoral species common for Europe and Mediterranean basin, and also by species widespread in the Palaearctic (92.9%). Autochthomous species constitute 5.5%, and species of the eastern Palaearctic origin, 1.6%. Hoverfly species with zoophagous larvae are the most widespread. The fraction of such species is low among species with narrowly local ranges (12.5%) and high among species with ranges including more than one dominion. Phytophagous species are wider represented among hoverflies with narrowly local ranges (44.7%) than among flies with interdominion ranges (8%). Ranges of dendrobiontic saprophagous species are determined by the presence of arboreal vegetation and no widespread polytopic hoverflies are found in this group. The distribution of mycetophagous species, commensals, and terrestrial saprophagous species is determined by the presence of fungi, ranges of some hymenopterans, and the presence of decaying plant remnants.     

The selective use of floral resources by the hoverfly Episyrphus balteatus (Diptera: Syrphidae) on farmland

The use of flowers of non-crop plants by the syrphid Episyrphus balteatus in margins of cereal fields was assessed quantitatively. G-tests and the ‘C’ index of Murdoch (1969) were used to compare the use and abundance of twenty-seven plant species by adult flies from June to August in the UK. The flowers of several plant species were used selectively by hoverflies throughout the season while the relative importance of others changed with date. The prospects for managing the field-margin flora to enhance biocontrol by hoverflies is discussed.     

Seasonal dynamics of hoverfly (Diptera,Syrphidae) activity in the taiga zone of the Komi Republic

194 species of 56 genera of hoverflies are recorded from the taiga zone of the Komi Republic. The phenology of the hoverfly flying activity and temporal structure of their trophic and ecological assemblages have been investigated. Ten phenological groups are distinguished and characteristics of adult diets of each of these groups are given.     

The conservation requirements of an endangered hoverfly, Hammerschmidtia ferruginea (Diptera, Syrphidae) in the British Isles

The saprophagous larva of the endangered hoverfly, Hammerschmidtia ferruginea (Fallén, 1817) (Diptera, Syrphidae) develops in wet, decaying sap under the bark of dead trees and branches of aspen, Populus tremula L. (Saliaceae). However this breeding site is transient: wet decay builds up patchily over 1.5–2 years of the tree or branch dying and lasts for a further 1–3 years before the bark falls off. Between 1990 and 2006, H. ferruginea swung through a cycle of abundance when the number of localities where it was detected dropped from 13 to 5 and back to 8. Fluctuations in amounts of dead wood caused by winds and storms probably explain this population swing. When there are few dead trees and branches, H. ferruginea breeds in sap flows on live P. tremula trees and populations are maintained but at low levels. To prevent local extinctions during periods when fallen wood is scarce, breeding habitat can be supplemented by felling trees and branches. Fallen wood with wet decaying sap is also important as an assembly site for mate selection. Adult food plants include flowers of bird cherry Prunus padus, rowan Sorbus aucuparia and hawthorn Crategus monogyna. Adults were recorded dispersing up to 1 km but are probably able to move further than this.     

Optical releasers of the innate proboscis extension in the hoverfly Eristalis tenax L. (Syrphidae,Diptera)

Freshly emerged, inexperienced imagos of the hoverfly Eristalis tenax L. extend their proboscis towards small, yellow colour stimuli, such as anther parts and artificial floral guides. The releasing of this behaviour, which is adapted to pollen feeding, was investigated in behavioural tests using white, UV-reflecting artificial flowers with four small screens illuminated with test stimuli serving as artificial floral guides. The releasing of the innate proboscis extension was tested using monochromatic test lights. Within an intensity range from approx. 5·10¹¹ to approx. 10¹⁴ quanta · cm^-2· s^-1, the flies extended their proboscis only towards green and yellow test lights (approx. 520–600 nm). The inhibition of the innate proboscis extension was tested using mixed light stimuli composed of a yellow monochromatic reference light (560 nm, 10¹³ quanta·cm^-2-1) and of a monochromatic test light. When the reference light was mixed with ultraviolet or blue test lights, the releasing of the innate proboscis extension was strongly inhibited, whereas admixing green/yellow light slightly promoted it; admixing red light had no effect. The results indicate that the releasing of the innate proboscis extension is mediated by the photoreceptor type R8y. Other receptor types which could cause the inhibition of the proboscis reaction are discussed.     

Floral preferences of hoverflies (Diptera: Syrphidae) in response to the abundance and species richness of flowering plants

For understanding the relationship between floral resources and the abundance of hoverflies, five meadows were selected at different elevations in a forested region. At each site the number of flowering herbaceous plants was assessed along a transect at 10-m intervals. To measure floral density, we counted the number of open flowers per species in each plot. Hoverflies visiting flowering plants were captured in each plot during 10 minutes. Path analysis showed that there was a negative correlation between plant species richness and syrphid abundance. Flower abundance had a strong positive direct effect on syrphid abundance but had indirect effects on the number of syrphid species. Only altitude had a strong positive direct effect on the number of syrphid species and this may be explained by the different climatic conditions. Some plant species attracted more syrphids than others. There were significantly fewer species of hoverfly per plot with increasing numbers of flowers. We interpret this to mean that plant species with a high density of flowers attracted more syrphids of the same species. Analysis of pollen grains extracted from two species of hoverflies showed that flower constancy occurred in three examined sites.     

Effect of hunger on yellow water trap catches of hoverfly (Diptera: Syrphidae) adults

1 An experiment was conducted in a winter wheat field using yellow water traps at crop height and at ground level, near to and distant from flowers, to test the hypothesis that such traps are seen as a source of food by flower‐feeding adult hoverflies and are therefore likely to selectively trap hungry individuals. Hoverflies caught in each trap were counted and identified and the amount of pollen in their guts was assessed. Ratios of numbers of hoverflies seen in the wheat crop to numbers caught in nearby traps were compared for the different treatments. 2 Most hoverflies were caught in crop‐high traps but they included a high proportion of individuals with empty guts. The taxa were: Episyrphus balteatus (De Geer), Metasyrphus corollae (F.) (these species together accounted for over 90% of the individuals trapped), with Sphaerophoria spp., Syrphus spp., Scaeva pyrastri (L.), Melanostomini and unidentified others making up the rest. 3 Significantly fewer hoverflies were captured in low than in high traps. The ratios of numbers trapped to numbers observed, in flower and in no flower treatments would be expected to be the same if the traps were not selective. This was supported for low traps. With high traps, however, there was a highly significant difference between the ratios (71.34 and 126 : 8, respectively). 4 Flies captured in high traps had less pollen in their gut than those captured in low ones. At each distance, more E. balteatus captured in high traps were in pollen category 1 (< 20 grains) than in any other category. The opposite state was seen in low trap catches, where most flies were in category 5 (> 5000 grains). Median pollen categories were 2 (21–200 grains) and 4 (1501–5000 grains) for flies caught in high and low traps, respectively. 5 The ecological selectivity of traps according to their height and the physiological condition of the targeted individuals is a problem likely to affect many trapping systems apart from the one described in this paper.     

Food utilization of aphidophagous hoverfly larvae (Diptera: Syrphidae, Chamaemyiidae) on herbaceous plants in an urban habitat

We examined food utilization in a community of aphidophagous hoverfly larvae (Diptera: Syrphidae and Chamaemyiidae) in open lands in an urban habitat in central Japan for 3 years. The community consisted of 17 hoverfly species feeding on 20 aphid species occurring on 14 species of dominant herbaceous plants. In terms of larval prey preference, the dominant eight species of hoverfly were categorized into three groups: a polyphagous ‘generalist’ group consisting of four species,Episyrphus balteatus, Betasyrphus serarius, Syrphus vitripennis andSphaerophoria sp.; an oligophagous ‘specialist’ group consisting of three species,Metasyrphus hakiensis, Dideoides latus andParagus hemorrhous; andLeucopis puncticornis, which showed a preference for two aphid species on the plantTorilis scabra. The prey aphids of the second group have behavioral or morphological defense mechanisms that are effective for preventing attacks by generalist hoverflies; two prey aphids are aggressive toward generalist predators and the others are protected by ant-attendance. The specialist hoverflies seem to be adapted to overcome these defense mechanisms. The prey ranges overlapped little between the generalist and the specialist groups, while those within the generalist group overlapped greatly.     

Field boundaries as barriers to movement of hover flies (Diptera: Syrphidae) in cultivated land

Field boundaries play an important role as refuges, food sources and corridors for invertebrates and vertebrates, and increasing farmland fragmentation impacts on these functions. However, hedgerows and other structures can also impede dispersal by flying insects. The current work uses the pollen of Phacelia tanacetifolia in hover fly guts as a marker to assess hover fly movement in farm landscapes. In the United Kingdom and New Zealand, Phacelia pollen was found in the guts of Ephisyrphus balteatus and Metasyrphus corollae (United Kingdom) and Melanostoma fasciatum (New Zealand) at distances up to 200 m from the source, when there were no barriers between the flowers and the traps used to catch the flies. The rate of decline over distance in the proportion of flies containing pollen was similar for the two countries. The extent to which four replicated field boundary types impeded hover fly movement was tested using post-and-wire fences, lines of poplars (Populus spp.) with gaps, dense poplars and controls (no potential barriers). Phacelia was planted on one side of each boundary, and along the centre of the control plots. The relative presence of the pollen in flies on both sides of the barriers showed that both types of poplar boundary restricted the movement of the flies, but the fence had no effect. In a separate experiment, gravid females of M. fasciatum were captured at a greater height on a shade-cloth fence than were non-gravid females and males. The implications of this work include the functioning and persistence of metapopulations and the influence of field boundaries on population recovery of beneficial invertebrates following pesticide-induced mortality. If field boundaries contribute to a temporal asynchrony between pest and natural enemy populations, this needs to be considered along with the well-established roles of boundaries as refugia for, and sources of, beneficial arthropods.     

Mark recapture estimates of dispersal ability and observations on the territorial behaviour of the rare hoverfly,Hammerschmidtia ferruginea (Diptera,Syrphidae)

In order to effectively manage habitat for fragmented populations, we need to know details of resource utilisation, and the capacity of species to colonise unoccupied habitat patches. Dispersal is vital in maintaining viable populations in increasingly fragmented environments by allowing re-colonisation of areas in which populations have gone extinct. In the UK, the endangered aspen hoverfly Hammerschmidtia ferruginea (Fallén 1817) (Diptera, Syrphidae) depends on a limited and transient breeding habitat: decaying aspen wood Populus tremula L. (Salicaceae). Conservation management for H. ferruginea involves encouraging aspen expansion across Scotland, and ensuring retention, maintenance and continuity of dead wood where H. ferruginea has been recorded and in areas that may link populations. In order to do this effectively we need to know how far H. ferruginea can disperse. By taking advantage of the tendency of adults to group on decaying aspen logs, we estimated dispersal ability through mark recapture techniques. In the first year, 1,066 flies were marked as they emerged from aspen logs and 78 were re-sighted at artificially-placed decaying aspen logs up to 4 km from the release site. In the second year, of 1,157 individually marked flies, 112 were re-sighted and one was observed 5 km from the release site. Territorial behaviour was recorded at all (19) decaying aspen log locations. In total, 72 males were recorded defending territories, which overlapped with 68 % of recorded female oviposition sites. Among males only, wing length was positively associated with dispersal. While these results show H. ferruginea is capable of locating decaying logs up to 5 km away, most dispersing individuals (68 %) were recorded at 1 km, which should be taken into account in developing management protocols. If enough dead wood is available it should be distributed within a radius of 1–2 km, and where possible, as stepping-stones linking up aspen woodlands. We discuss the implications of our findings for the natural history of this species, and make recommendations for its conservation management.     

Diurnal activity patterns in hoverflies (Diptera, Syrphidae)

Abstract. 1. Time-budgets are constructed from censuses of hoverflies. Larger species spend less time in flight.
2. Most species are active under similar conditions of light intensity, but a shade-tolerant (Melanostoma scalare Fabr.) and a sun-loving species (Metasyrphus corollae Fabr.) were identified.
3. Large species become active (i.e. move, by flying or feeding) at lower temperatures than small ones, except M.scalare, which is tolerant to cool temperatures. This means that, M.scalare apart, the order of species appearing during the day is size-dependent, largest first.
4. Previous claims about the diel periodicity of syrphids can be reconciled by considering the importance of individual thermal balance.
5. Tolerance to low temperatures in small hoverflies is associated with feeding on anemophilous pollen; this may indicate a proline-fuelled flight.     

Functional richness, functional evenness, and use of niche space in parasite communities

Most efforts aimed at elucidating the factors responsible for the variation in species richness among different parasite communities have focused on host characteristics such as body mass or diet. Independently of host features, however, the way in which parasites use resources within the host may also affect the species richness of the community. The distribution of parasite individuals or biomass in niche space determines whether host resources are used evenly, or whether there are gaps in resource use, with some parts of the total niche being underutilized. Here, the concepts of functional richness and functional evenness are applied to parasite communities for the first time, using simple indices. Measurements of the distribution of species in niche space within communities, such as mapping the distribution of helminths along the length of the host's intestine, is standard practice in parasitology. In such cases, functional richness is simply the proportion of the total number of intestine sections available that are used by at least 1 worm, whereas functional evenness measures the evenness in the distribution of worm numbers or biomass across all niche sections that are occupied. Data on cestode communities of elasmobranchs are used to illustrate the use of these indices, and to show how important they can be in tests of ecological hypotheses. The indices presented here capture essential features of resource use in parasite communities, and can be useful tools for comparative analyses.     

Blackfly (Diptera, Simuliidae) communities and species richness estimation in Carpathian montane streams

The present work gives the result of blackfly fauna research of the Slovenský raj (West Carpathians, Slovakia). Our aim was to describe the composition of blackfly communities, find indicator species for communities, describe environmental factors affecting communities and determine the potential changes in species richness. Research was carried out in 2007 and 2008. The presence of 22 species of blackflies was confirmed at 20 sites in the area of Slovenský raj. 12 species were recorded for the first time in this area. The potential for further increase in diversity was calculated by a nonparametric estimation of species richness. Two main groups of sites with their typical blackflies communities were indentified using a TWINSPAN analysis. The presence of Simulium ornatum, S. variegatum and S. argyreatum was typical in larger streams in broad valleys. The absence or low abundance of these species was typical for smaller streams in canyons. Based on the species composition of blackflies communities, we identified 6 groups of sites: 1. anthropogenically disturbed submountain rivers (typical species: Simulium ornatum, S. reptans, S. equinum), 2. undisturbed metarhithral (typical species: Simulium bertrandi, S. codreanui, S. costatum, S. vernum, S. argenteostriatum, S. degrangei, S. argyreatum, S. variegatum), 3. undisturbed hyporhithral (typical species: Simulium trifasciatum, S. tuberosum), 4. undisturbed epirhithral (typical species: Prosimulium hirtipes, P. rufipes), 5. hypocrenal (typical species: Simulium cryophilum, S. monticola) and 6. anthropogenically disturbed epirhithral (typical species: Simulium brevidens, S. maximum). The most important environmental factors explaining differences in species composition like altitude, flow speed, pH, conductivity, oxygen saturation, sulphates concentration and presence of riparian vegetation were identified by the CCA analysis.     

Functional traits as indicators of biodiversity response to land use changes across ecosystems and organisms

Rigorous and widely applicable indicators of biodiversity are needed to monitor the responses of ecosystems to global change and design effective conservation schemes. Among the potential indicators of biodiversity, those based on the functional traits of species and communities are interesting because they can be generalized to similar habitats and can be assessed by relatively rapid field assessment across eco-regions. Functional traits, however, have as yet been rarely considered in current common monitoring schemes. Moreover, standardized procedures of trait measurement and analyses have almost exclusively been developed for plants but different approaches have been used for different groups of organisms. Here we review approaches using functional traits as biodiversity indicators focussing not on plants as usual but particularly on animal groups that are commonly considered in different biodiversity monitoring schemes (benthic invertebrates, collembolans, above ground insects and birds). Further, we introduce a new framework based on functional traits indices and illustrate it using case studies where the traits of these organisms can help monitoring the response of biodiversity to different land use change drivers. We propose and test standard procedures to integrate different components of functional traits into biodiversity monitoring schemes across trophic levels and disciplines. We suggest that the development of indicators using functional traits could complement, rather than replace, the existent biodiversity monitoring. In this way, the comparison of the effect of land use changes on biodiversity is facilitated and is expected to positively influence conservation management practices.     

黑带食蚜蝇Episyrphus balteatus De Geer的复眼结构及其调光机制

【目的】观察研究黑带食蚜蝇Episyrphus balteatus De Geer成虫复眼形态、小眼结构和不同光暗条件对小眼结构的影响,以明确其光视觉的结构基础和调光机制。【方法】利用组织切片法和扫描电镜等技术。【结果】1.复眼位于头部两侧,正面观呈半球形,占据除额颜外大部分头部。雄虫与雌虫单个复眼分别有约7 180个、7 230个小眼。各小眼面呈整齐排列的规则六边形。2.小眼由角膜及伪晶锥组成的屈光器、不同水平面分布的8个小网膜细胞及其特化形成的离散型视杆、屏蔽色素细胞和基膜等组成。小眼自远端至近端由主色素细胞和12个附属色素细胞围绕。3.随光暗条件的改变小眼内的附属色素细胞色素和基细胞细胞核沿小眼纵轴移动。光适应时,附属色素细胞色素颗粒沿小眼纵轴均匀分布,基细胞细胞核位于基膜上方。暗适应时,附属色素细胞色素颗粒向伪晶锥近端压缩,基细胞细胞核亦向远端移动,到达视杆中段。【结论】黑带食蚜蝇复眼精密的小眼排列形式和内部结构均显示了其强大的生理功能;屏蔽色素颗粒的移动是其复眼适应外界光环境变化的重要机制。本试验为进一步探究黑带食蚜蝇视觉结构和光调节机制,以及与其飞行行为间的关系提供了一定的理论基础。     

Agricultural land use and black fly (Diptera,Simuliidae) species richness and species assemblages in tropical streams,Northeastern Thailand

Habitat degradation through agricultural land use is the major factor threatening lotic ecosystems. Although black flies are major components of these ecosystems, the impact of agricultural land use on species diversity and species assemblages has been largely ignored in tropical streams of the Oriental region. The objectives of this study are to examine patterns of species distribution and species richness and to compare black fly species richness and species assemblages in forest and agricultural streams in Thailand. A total of 143 collections were made from 70 stream sites between June 2007 and May 2008. Whereas 19 black fly species found in these collections were all found in forest sites, only 13 species were found in agricultural sites. High species richness was associated with larger, faster, and cooler streams with larger streambed particles and the presence of riparian trees. Logistic regression analyses revealed that stream size, velocity, and riparian vegetation are among the most important factors determining patterns of spatial distribution. The results are largely consistent with studies in other zoogeographic regions, suggesting the existence of general rules for black fly species distributions. Comparisons of the physicochemical conditions between forest and agricultural streams indicated that streams in agricultural areas are warmer, with higher conductivity and fewer riparian trees. Species richness was significantly higher in forest than in agricultural streams (t = 3.61, P < 0.001). Streams in forest areas were predominantly occupied by S. siamense (73%) but other species were also found at a relatively high frequency (>20%) of the sampling sites. In contrast, streams in agricultural areas were predominantly occupied by S. aureohirtum (>80%) among the sole black fly species at 27% of the sites. The results indicate that agricultural land use has a significantly detrimental impact on black fly diversity and species assemblages. Handling editor: D. Dudgeon     

Assessing the effects of vegetation type on hoverfly (Diptera: Syrphidae) diversity in a Mediterranean landscape: implications for conservation

Understanding the responses of insects to ecological variables provides information that is fundamental for their conservation. The present study took place in three different landscapes (three plots of 10 × 10 km each) in a typical Mediterranean ecosystem of a Spanish national park. Each landscape included three vegetation types, grasslands, scrublands, and woodlands, and was characterised by a dominant vegetation type. Our objectives were: (1) to assess how important the influence of the dominant vegetation type and the vegetation type of sites are on hoverfly (Diptera: Syrphidae) diversity at landscape scale; (2) to assess whether scrublands are contributing to the loss of hoverfly diversity in an ecosystem with a long history of human use. In order to achieve these goals, we compared hoverfly diversity among sampling sites by prospecting all three diversity levels, alpha, beta and gamma, at each landscape. We sampled adult hoverflies at 18 sites located in different vegetation types within the three landscapes. No evidence was found that demonstrated an effect of the dominant vegetation type on alpha or beta diversity of sites, but the vegetation type of sites did have an affect. The highest species richness was found in woodland sites, particularly in the grassland-dominated landscape. At each landscape, beta diversity among sampling sites contributed more to gamma diversity than alpha diversity did. Our results highlight the need to focus on the conservation of woodland remnants of grassland-dominated landscape and scrubland-dominated landscape in order to preserve a large proportion of the biodiversity of Cabañeros hoverflies, as well as on the maintenance of the mosaic landscape, which is linked to high beta diversity, typical in many Mediterranean ecosystems. We emphasise the importance of open clearings in the vast mass of scrubland in the scrubland-dominated landscape in order to provide extra resources for the hoverflies.     

Internal labelling of the eggs of the aphidophagous hoverfly Episyrphus balteatus (De Geer) (Diptera: Syrphidae) with a fluorescent dye

A method was developed to label the egg load of Episyrphus balteatus (De Geer) (Diptera: Syrphidae) by feeding adult females with the fluorescent dye Rhodamine B. In a preliminary experiment aimed at selecting an appropriate dye concentration, gravid females reared in insectary conditions were provided for two successive days with a 50% honey-water solution enriched with Rhodamine B at concentrations of 0.1%, 0.01% and 0.001%. Eggs deposited on aphid-infested broad bean plants by syrphid females fed with Rhodamine B 0.01% showed the most intense marking. The higher dye concentration induced irregular marking among the eggs, presumably due to its repellent taste and consequently lower intake by the flies. A second series of trials focused on the persistence of the marker, prior and after oviposition, at the selected (optimal) dilution. A reduction in the marking degree of eggs was observed when the first egg deposition was delayed for one day after removing the dye from the cage. When marked females were allowed to deposit further batches of eggs the subsequent days, fewer eggs were laid and their fluorescence properties was reduced. Exposure to direct sunlight during 10 hours had no detectable effect on the fluorescence level of marked eggs deposited on a bean plant. The egg-marking technique examined in this study could be useful to determine the crop area accommodating eggs deposited by marked syrphid females migrating from a release point into cultivated fields.     

Trophic links,nutrient fluxes,and natural history in the Allium ursinum food web,with particular reference to life history traits of two hoverfly herbivores (Diptera: Syrphidae)

The food web centering on Allium ursinum (Liliaceae) in a beech forest (Germany) is described, and temporal variation of active trophic links is related to species' life cycles. The most important insect herbivores are Cheilosia fasciata (a larval leaf miner) and Portevinia maculata (a larva bulb miner) (Diptera: Syrphidae). Energy, carbon and nitrogen flow in the food chain (Allium-Cheilosia-Phygadeuon ursini) are investigated and analysed with respect to differences in resource allocation by the leaf miner and its hymenoptereous parasitoid. In C. fasciata nitrogen is likely to be the limiting resource, while growth in Phygadeuon ursini appears energy-limited. Larval feeding habits of C. fasciata and Portevinia maculata determined the timing of the species' life cycles and, as a consequence, appeared to preclude the existence of a pupal parasitoid in Portevinia maculata. Further details of life history traits are demonstrated and discussed.