Caught in the web: Spider web architecture affects prey specialization and spider–prey stoichiometric relationships

Quantitative approaches to predator–prey interactions are central to understanding the structure of food webs and their dynamics. Different predatory strategies may influence the occurrence and strength of trophic interactions likely affecting the rates and magnitudes of energy and nutrient transfer between trophic levels and stoichiometry of predator–prey interactions. Here, we used spider–prey interactions as a model system to investigate whether different spider web architectures—orb, tangle, and sheet‐tangle—affect the composition and diet breadth of spiders and whether these, in turn, influence stoichiometric relationships between spiders and their prey. Our results showed that web architecture partially affects the richness and composition of the prey captured by spiders. Tangle‐web spiders were specialists, capturing a restricted subset of the prey community (primarily Diptera), whereas orb and sheet‐tangle web spiders were generalists, capturing a broader range of prey types. We also observed elemental imbalances between spiders and their prey. In general, spiders had higher requirements for both nitrogen (N) and phosphorus (P) than those provided by their prey even after accounting for prey biomass. Larger P imbalances for tangle‐web spiders than for orb and sheet‐tangle web spiders suggest that trophic specialization may impose strong elemental constraints for these predators unless they display behavioral or physiological mechanisms to cope with nutrient limitation. Our findings suggest that integrating quantitative analysis of species interactions with elemental stoichiometry can help to better understand the occurrence of stoichiometric imbalances in predator–prey interactions.     

Diet-dependent fecundity of the spiders Atypena formosana and Pardosa pseudoannulata, predators in irrigated rice

1 The fecundity of the spiders Atypena formosana and Pardosa pseudoannulata was assessed on diets of brown planthopper Nilaparvata lugens (BPH), green leafhopper Nephotettix virescens (GLH), Collembola (Entomobryidae), Droso phila melanogaster and three prey mixtures; BPH–GLH, BPH–GLH–Collembola and a full mixed diet of BPH–GLH–Collembola–D. melanogaster. 2 Egg production and hatching success was high in A. formosana except on the BPH, the GLH and the BPH–GLH diets. In P. pseudoannulata egg production and hatching success was lowest on diets of GLH and BPH–GLH, whereas this spider used BPH better. 3 Differences between offspring sizes were small. 4 Survival of A. formosana females was lowest on the GLH and BPH–GLH diets. In P. pseudoannulata survival did not differ significantly. 5 Food conversion efficiency was highest on the Collembola and the D. melanogaster diets for both spiders. 6 Overall, diets of Collembola and D. melanogaster had the highest quality for both spiders, the BPH–GLH–Collembola and the full mixed diets were intermediate in quality, the quality of the BPH diet was intermediate to low and quality of the GLH and BPH–GLH diets was low. 7 There need not be a contradiction between low dietary value of hoppers and successful natural biological control provided that the nutritional needs of the spiders are met by sufficient alternative prey. 8 The hopper‐spider relationship is comparable to that of aphids and spiders in temperate cereals, suggesting that low dietary quality of Homopterans to spiders may be widespread.     

Selectivity underlies the dissociation between seasonal prey availability and prey consumption in a generalist predator

Generalist predators are capable of selective foraging, but are predicted to feed in close proportion to prey availability to maximize energetic intake especially when overall prey availability is low. By extension, they are also expected to feed in a more frequency‐dependent manner during winter compared to the more favourable foraging conditions during spring, summer and fall seasons. For 18 months, we observed the foraging patterns of forest‐dwelling wolf spiders from the genus Schizocosa (Araneae: Lycosidae) using PCR‐based gut‐content analysis and simultaneously monitored the activity densities of two common prey: springtails (Collembola) and flies (Diptera). Rates of prey detection within spider guts relative to rates of prey collected in traps were estimated using Roualdes’ c_st model and compared using various linear contrasts to make inferences pertaining to seasonal prey selectivity. Results indicated spiders foraged selectively over the course of the study, contrary to predictions derived from optimal foraging theory. Even during winter, with overall low prey densities, the relative rates of predation compared to available prey differed significantly over time and by prey group. Moreover, these spiders appeared to diversify their diets; the least abundant prey group was consistently overrepresented in the diet within a given season. We suggest that foraging in generalist predators is not necessarily restricted to frequency dependency during winter. In fact, foraging motives other than energy maximization, such as a more nutrient‐focused strategy, may also be optimal for generalist predators during prey‐scarce winters.     

A specialized araneophagic predator's short‐term nutrient utilization depends on the macronutrient content of prey rather than on prey taxonomic affiliation

A specialist predator that has a specialized diet, prey‐specific prey‐capture behaviour and a preference for a particular type of prey may or may not be specialized metabolically. Previous studies have shown that jumping spiders of the genus Portia prey on other spiders using prey‐specific prey‐capture behaviour, prefer spiders as prey to insects and gain long‐term benefits in terms of higher survival and growth rates on spider diets than on insect diets. However, it is unclear whether there are substances uniquely present in spiders on which Portia depends, or, alternatively, spiders and insects all contain more or less the same nutrients but the relative amounts of these substances are such that Portia perform better on a spider diet. These questions are addressed by testing the hypothesis that prey specialization includes metabolic adaptations that allow Portia an enhanced nutrient extraction or nutrient utilization efficiency when feeding on spider prey compared with insect prey. Three groups of Portia quei Zabka are fed either their preferred spider prey or one of two types of flies (Drosophila melanogaster Meigen) that differ in nitrogen and lipid content. Portia quei shows a higher feeding rate of high‐protein flies than of high‐lipid flies and spiders but, after 5 days of feeding, there is no significant difference in growth between treatments, and the diets lead to significant changes in the macronutrient composition of P. quei as a result of variable extraction and utilization of the prey. The short‐term utilization of spider prey is similar to that of high‐lipid flies and both differ in several respects from the utilization of high‐protein flies. Thus, the short‐term nutrient utilization is better explained by prey macronutrient content than by whether the prey is a spider or not. The results suggest that spider prey may have a more optimal macronutrient composition for P. quei and that P. quei does not depend on spider‐specific substances.     

Body size and tree species composition determine variation in prey consumption in a forest‐inhabiting generalist predator

Trophic interactions may strongly depend on body size and environmental variation, but this prediction has been seldom tested in nature. Many spiders are generalist predators that use webs to intercept flying prey. The size and mesh of orb webs increases with spider size, allowing a more efficient predation on larger prey. We studied to this extent the orb‐weaving spider Araneus diadematus inhabiting forest fragments differing in edge distance, tree diversity, and tree species. These environmental variables are known to correlate with insect composition, richness, and abundance. We anticipated these forest characteristics to be a principle driver of prey consumption. We additionally hypothesized them to impact spider size at maturity and expect shifts toward larger prey size distributions in larger individuals independently from the environmental context. We quantified spider diet by means of metabarcoding of nearly 1,000 A. diadematus from a total of 53 forest plots. This approach allowed a massive screening of consumption dynamics in nature, though at the cost of identifying the exact prey identity, as well as their abundance and putative intraspecific variation. Our study confirmed A. diadematus as a generalist predator, with more than 300 prey ZOTUs detected in total. At the individual level, we found large spiders to consume fewer different species, but adding larger species to their diet. Tree species composition affected both prey species richness and size in the spider''s diet, although tree diversity per se had no influence on the consumed prey. Edges had an indirect effect on the spider diet as spiders closer to the forest edge were larger and therefore consumed larger prey. We conclude that both intraspecific size variation and tree species composition shape the consumed prey of this generalist predator.     

Are you what you eat? A highly transient and prey‐influenced gut microbiome in the grey house spider Badumna longinqua

Stable core microbial communities have been described in numerous animal species and are commonly associated with fitness benefits for their hosts. Recent research, however, highlights examples of species whose microbiota are transient and environmentally derived. Here, we test the effect of diet on gut microbial community assembly in the spider Badumna longinqua. Using 16S rRNA gene amplicon sequencing combined with quantitative PCR, we analyzed diversity and abundance of the spider's gut microbes, and simultaneously characterized its prey communities using nuclear rRNA markers. We found a clear correlation between community similarity of the spider's insect prey and gut microbial DNA, suggesting that microbiome assembly is primarily diet‐driven. This assumption is supported by a feeding experiment, in which two types of prey—crickets and fruit flies—both substantially altered microbial diversity and community similarity between spiders, but did so in different ways. After cricket consumption, numerous cricket‐derived microbes appeared in the spider's gut, resulting in a rapid homogenization of microbial communities among spiders. In contrast, few prey‐associated bacteria were detected after consumption of fruit flies; instead, the microbial community was remodelled by environmentally sourced microbes, or abundance shifts of rare taxa in the spider's gut. The reshaping of the microbiota by both prey taxa mimicked a stable core microbiome in the spiders for several weeks post feeding. Our results suggest that the spider's gut microbiome undergoes pronounced temporal fluctuations, that its assembly is dictated by the consumed prey, and that different prey taxa may remodel the microbiota in drastically different ways.     

Collembola as alternative prey sustaining spiders in arable ecosystems: prey detection within predators using molecular markers

Collembola comprise a major source of alternative prey to linyphiid spiders in arable fields, helping to sustain and retain these predators as aphid control agents within the crop. Polymerase chain reaction primers were developed for the amplification, from spider gut samples, of DNA from three of the most abundant species of Collembola in wheat crops in Europe, namely Isotoma anglicana, Lepidocyrtus cyaneus and Entomobrya multifasciata. The primers amplified fragments of the mitochondrial cytochrome oxidase subunit I (COI) gene and were designed following alignment of comparable sequences for a range of predator and prey species. Each of the primer pairs proved to be species-specific to a Collembola species, amplifying DNA fragments from 211 to 276 base pairs in length. Following consumption of a single collembolan, prey DNA was detectable in 100% of spiders after 24 h of digestion. We report the first use of DNA-based techniques to detect predation by arthropods on natural populations of prey in the field. All three species of Collembola were consumed by the spiders. By comparing the ratios of the Collembola species in the field with the numbers of spiders that gave positive results for each of those species, it was possible to demonstrate that the spiders were exercising prey choice. Overall, a single target species of Collembola was eaten by 48% of spiders while a further 16% of spiders contained DNA from two different species of Collembola. Preference was particularly evident for I. anglicana, the species most frequently found in spider guts yet the least numerous of the three target species in the field.     

The value of two Collembola species as food for a linyphiid spider

Egg production by the cereal spider Erigone atra was used as a fitness parameter for evaluating the food quality of two species of Collembola: Folsomia fimetaria and Isotoma anglicana. Drosophila melanogaster was used as reference prey. We tested the hypothesis that due to differences in food quality, the two Collembola species would affect the reproduction of the spider differently. The quality ranking of the prey types turned out as: I. anglicana > D. melanogaster > F. fimetaria. With F. fimetaria alone, spiders were unable to maintain reproduction. E. atra was more efficient in utilising I. anglicana and D. melanogaster. Thus, daily consumption rates of I. anglicana were lower in spite of higher egg laying rates by E. atra. A mixed diet of F. fimetaria and D. melanogaster resulted in a lower reproductive output than a pure diet of D. melanogaster, indicating a toxic element in F. fimetaria. In the mixed diet F. fimetaria had a negative influence on the consumption capacity of the spider towards D. melanogaster, while D. melanogaster had a positive influence on the consumption capacity towards F. fimetaria. It is concluded that a high abundance of I. anglicana may support a high reproductive output of E. atra, while the presence, of F. fimetaria in fields may reduce the spider's reproductive output.     

Evolution of stenophagy in spiders (Araneae): evidence based on the comparative analysis of spider diets

Stenophagy (narrow diet breadth) represents an extreme of trophic specialization in carnivores, but little is known about the forces driving its evolution. We used spiders, the most diversified group of terrestrial predators, to investigate whether stenophagy (1) promoted diversification; (2) was phylogenetically conserved and evolutionarily derived state; and (3) was determined either by geographical distribution and foraging guild. We used published data on the prey of almost 600 species. Six categories of stenophagy were found: myrmecophagy, araneophagy, lepidopterophagy, termitophagy, dipterophagy, and crustaceophagy. We found that the species diversity of euryphagous genera and families was similar to stenophagous genera and families. At the family level, stenophagy evolved repeatedly and independently. Within families, the basal condition was oligophagy or euryphagy. Most types of stenophagy were clearly derived: myrmecophagy in Zodariidae; lepidopterophagy in Araneidae; dipterophagy in Theridiidae. In contrast, araneophagy was confined to basal and intermediate lineages, suggesting its ancestral condition. The diet breadth of species from the tropics and subtropics was less diverse than species from the temperate zone. Diet breadth was lower in cursorial spiders compared to web-building species. Thus, the evolution of stenophagy in spiders appears to be complex and governed by phylogeny as well as by ecological determinants.     

Invertebrate biodiversity affects predator fitness and hence potential to control pests in crops

Natural enemies that control pests usually allow farmers to avoid, or reduce, the use of pesticides. However, modern farming practices, that maximize yields, are resulting in loss of biodiversity, particularly prey diversity. Does this matter? Pests continue to thrive, and without alternative prey the predators should, perforce, concentrate their attentions upon the pests.We showed that a diverse diet significantly enhances predator fecundity and survival. Experiments were conducted using common generalist predators found in arable fields in Europe, the carabid beetle Pterostichus melanarius (Coleoptera: Carabidae) and the linyphiid spider Erigone atra (Araneae: Linyphiidae). We tested the hypothesis that mixed species diets were optimal, compared with restricted diets, with respect to parameters such as predator weights, egg weights, numbers of eggs laid, egg development times, egg hatching rates and predator survival. In carabids, an exclusive earthworm diet was as good as mixed diets containing earthworms for egg production and hatching, but less good than such mixed diets for increase in beetle mass and sustained egg laying. For spiders, aphids alone (Sitobion avenae) or with the Collembola Folsomia candida, drastically reduced survival. Aphids plus the Collembola Isotoma anglicana improved survival but only aphids with a mixed Collembola diet maximized numbers of hatching eggs.Predators offered only pests (slugs or aphids) had lowest growth rates and fecundity. We therefore demonstrated that conservation of a diversity of prey species within farmland, allowing predators to exploit a diverse diet, is essential if predators are to continue to thrive in crops and regulate agricultural pests.     

Prey preferences ofPortia fimbriata,an araneophagic,web-building jumping spider (Araneae: Salticidae) from Queensland

Portia fimbriata from Queensland, a previously studied jumping spider (Salticidae), routinely includes web-building spiders and cursorial salticids in its diet, both of these types of prey being dangerous and unusual prey for a salticid. The present paper is the first detailed study ofP. fimbriata's prey preferences. Three basic types of tests of prey preference were used, providing evidence that (1)P. fimbriata males and females prefer spiders (both web-building spiders in webs and salticids away from webs) to insects; (2)P. fimbriata males and females prefer salticids to web-building spiders; (3)P. fimbriata males and females prefer larger spiders to smaller spiders; (4) there are intersexual differences in the preferences ofP. fimbriata for prey size, females preferring larger prey and males preferring smaller prey; and (5)P. fimbriata's prey preferences are not affected by a prior period without food of 2 weeks. When preferences were tested for by using both living, active prey and dead, motionless lures, the same preferences were expressed, indicating thatP. fimbriata can distinguish among different types of prey independent of the different movement patterns of different prey.     

More than a barrier: The complex effects of ecotone vegetation type on terrestrial consumer consumption of an aquatic prey resource

Community structure and dynamics can be influenced by resource transfers between ecosystems, yet little is known about how boundary structure determines both the magnitude of exchanges and their effects on recipient and donor communities. Aquatic and terrestrial ecosystems are often linked by resource fluxes and riparian vegetation is commonly affected by anthropogenic alterations to land use or river hydrological regime. I investigated whether shrubs at the freshwater–terrestrial interface alter the supply, distribution and importance of aquatic prey resources to terrestrial consumers. Shrubs were predicted to alter the larval community composition of aquatic insects and the emergence of winged adults, thus affecting aquatic prey subsidies to terrestrial consumers. In addition, shrubs were hypothesized to alter the microclimatic suitability of the riparian zone for adult aquatic insects, act as a physical barrier to their dispersal and affect terrestrial community composition, particularly the abundance and type of predators that could benefit from the aquatic prey resource. Stable isotope dietary analyses and a survey of shrub‐dominated and open grassland riparian habitats revealed that larval densities of aquatic insects (EPTM: Ephemeroptera, Plecoptera, Trichoptera and Megaloptera) were higher in shrub than grassland habitats; however, reduced emergence and lateral dispersal in shrub areas led to lower densities of adults. The temperature and relative humidity of the riparian zone did not differ between the habitats. Ground‐active terrestrial invertebrate communities had a higher proportion of cursorial spiders in grassland, coinciding with greater abundances of aquatic prey. Aquatic prey contribution to cursorial spider diet matched adult aquatic insect abundances. Overall, riparian shrubs reduced the magnitude, or at least altered the timing, of cross‐ecosystem subsidy supply, distribution and use by consumers through mechanisms operating in both the aquatic and terrestrial ecosystems. Thus, the structure of ecosystem boundaries has complex effects on the strength of biological interactions between adjacent systems.     

Density‐dependent and ‐independent effects on the joint use of space by predators and prey in terrestrial arthropod food‐webs

The spatial distribution of predators and their prey is affected by their joint use of space. While the formation of such spatial patterns may be driven by density‐dependent and ‐independent factors our knowledge on the contribution of different land‐use activities on the formation of spatial patterns between predators and prey remains very limited. Agriculture is one of the most prevailing land‐use activities with strong effects on invertebrate densities and structural habitat conditions. Here, we used replicated conventionally and organically managed winter wheat fields to investigate the effects of agricultural land‐use on the spatial patterns of generalist predators and decomposer prey. We then identified the explanatory power of density‐dependent (prey and predator activity density) and density‐independent (vegetation structure) predictors for the observed spatial patterns. Generalist predators were regularly distributed only in conventionally managed fields and this pattern intensified with decreasing Collembola prey availability and increasing spider activity density. Segregation between carabid and spider predators was strongest in fields with lowest wheat plant height, suggesting more intense intraguild interactions in structurally less complex habitats. Collembola were aggregated independent of management and aggregation was strongest in fields with highest Collembola and carabid activity density. Spiders and Collembola prey were associated, but higher aphid densities under conventional management weakened or interrupted this spatial relationship. We conclude that active control of crop plant physiognomy by growth hormones and herbicides in conventionally managed fields promotes predator–predator segregation and that a high availability of aphid prey seems to decouple predator–Collembola prey associations. Our results emphasise the need for a more mechanistic understanding of the effects of land‐use on the formation of spatial patterns and species interactions, especially under scenarios of environmental change and an ongoing loss of biodiversity.     

Crab spider hunting performance is temperature insensitive

1. Laboratory and field experiments showed that the hunting performance of two flower-dwelling crab spiders, Misumenops asperatus and Misumenoides formosipes, was thermally insensitive over a broad range of temperatures normally experienced by these spiders. 2. In the laboratory, HP, a behavioural metric of spider hunting performance, was similar for spiders of a given species over an ≈ 30 °C temperature range. 3. Spiders in the field captured predominantly hymenopterans and dipterans, and field hunting performance, measured as the number of prey captured per spider per day, also proved to be unaffected by temperature. 4. These findings counter the general rule that physiological/ecological performance in terrestrial arthropods is temperature dependent. 5. Freedom from temperature constraints on the capacity of crab spiders to capture prey may be due to the use of venom and/or to muscle physiological adaptations for anaerobic metabolism. 6. Wide thermal performance breadth increases the spectrum of prey available to M. asperatus and M. formosipes by allowing spiders to hunt prey active during cooler periods of the day as well as those active during warmer periods. 7. Wide thermal performance breadth also benefits M. asperatus and M. formosipes due to adult phenology; both species experience a seasonal temperature shift during the adult phase.     

The effect of colour variation in predators on the behaviour of pollinators: Australian crab spiders and native bees

1. Australian crab spiders exploit the plant–pollinator mutualism by reflecting UV light that attracts pollinators to the flowers where they sit. However, spider UV reflection seems to vary broadly within and between individuals and species, and we are still lacking any comparative studies of prey and/or predator behaviour towards spider colour variation. 2. Here we looked at the natural variation in the coloration of two species of Australian crab spiders, Thomisus spectabilis and Diaea evanida, collected from the field. Furthermore, we examined how two species of native bees responded to variation in colour contrast generated by spiders sitting in flowers compared with vacant flowers. We used data from a bee choice experiment with D. evanida spiders and Trigona carbonaria bees and also published data on T. spectabilis spiders and Austroplebeia australis bees. 3. In the field both spider species were always achromatically (from a distance) undetectable but chromatically (at closer range) detectable for bees. Experimentally, we showed species‐specific differences in bee behaviour towards particular spider colour variation: T. carbonaria bees did not show any preference for any colour contrasts generated by D. evanida spiders but A. australis bees were more likely to reject flowers with more contrasting T. spectabilis spiders. 4. Our study suggests that some of the spider colour variation that we encounter in the field may be partly explained by the spider's ability to adjust the reflectance properties of its colour relative to the behaviour of the species of prey available.     

Multichannel feeding by spider functional groups is driven by feeding strategies and resource availability

Multichannel feeding, whereby consumers feed across resource channels such as upon herbivore and detritivore resources, acts to link discrete compartments of a food web with implications for ecosystem functioning and stability. Currently however, we have little understanding which feeding strategies of consumers underlie multichannel feeding. We therefore link spider functional group and resource density‐dependent or density‐independent feeding strategies to multichannel feeding by quantifying not only consumer diet, but also the relative availability of resources. Here we analysed herbivore (green) and detritivore (brown) prey use by spider communities in grasslands, and tested if available prey biomass proportions were linked to observed spider dietary proportions. Different spider functional groups each linked green and brown resource channels, but while green prey were always consumed in proportion to their relative biomass, brown prey were consumed independently of proportion by some functional groups. Additionally, we found greater intraguild predation by cursorial spiders when green resources were relatively scarcer, suggesting green prey was preferred, and needed to be compensated for when rare. Overall, we observed a stronger consumer connection to the green than brown resource channel, yet this green connection was more variable due to greater range in green resource availability across grasslands and density‐dependent consumption on green prey. Consequently, multichannel feeding by spiders was determined by density‐dependent and density‐independent feeding strategies that varied by spider functional group and across resources channels. Our results demonstrate that the role of multichannel feeding by spiders in linking separate food web compartments is a dynamic component of food web structure in these wild grasslands.     

Differences in venom composition between orb-weaving and wandering Hawaiian Tetragnatha (Araneae)

Spider venoms are complex mixtures of toxins that are used primarily for immobilizing prey. There is evidence of chemical variation in spider venoms among close relatives, yet few studies have analysed their evolution within an ecological and phylogenetic framework. On the Hawaiian archipelago, Tetragnatha, a cosmopolitan orb-weaving genus, has undergone a radiation in which a monophyletic lineage has abandoned web-building and become obligately wandering foragers. This study compares venom composition and details of feeding behaviour between orb-weaving and wandering Hawaiian Tetragnatha. Protein gel electrophoresis patterns indicated that relative to orb-weavers, wandering species had a reduced concentration of low molecular weight (<14kDa) components. Both orb-weaving and wandering Tetragnatha captured flying prey (adult lepidopterans, dipterans), but wandering spiders also captured non-flying prey (insect larvae, spiders). There were no distinct differences between orb-weavers and wanderers in prey capture and immobilization sequences, or in the paralytic effects of bites on prey. However, prey bitten by wanderers took longer to be permanently immobilized than prey bitten by orb-weavers. Contrary to predictions, there was no indication that web-loss in this group was associated with an increase in venom potency.     

The time scale of isotope signals in spiders: molting the remains of a previous diet

Stable isotope analysis (SIA) has emerged as an important tool for understanding consumer diets and diet shifts. However, although the general idea behind SIA is clear, the interpretation of data is often fraught with problems because tissue turnover and fractionations are not known. We investigated shifts in stable isotope composition of spiders following a diet shift, using mealworms fed either maize (C4) or wheat (C3) flour. Mealworms had different carbon isotope composition depending on their diet and this difference was reflected in spider body parts. In the experiment, we first fed the spiders on a diet of either maize‐fed or wheat‐fed mealworms and then switched diet at the time of the second molt. Spiders were then sampled repeatedly until the next molt. We sampled both legs and abdomens, as these are presumed to have different turnover of tissue, and also molt remains were sampled when this was relevant. The data indicated that the spider legs had a turnover of about 20 days, whereas the spider abdomens had a turnover of about 8 days. Molt remains had the slowest turnover and reflected the diet at the previous molt, when the exoskeleton was formed. Both these observations indicate that SIA may be successfully used for elucidating diet shifts. More problematic was the fact that fractionation of carbon isotope ratios varied with body parts and diets. When spiders were fed maize‐mealworms then the fractionation was larger for abdomens, but when the spiders were fed wheat‐mealworms then the fractionation was larger for legs. The mechanisms underlying this pattern are unclear and deserve further attention.     

Cursorial spiders and succession: age or habitat structure?

Summary The structure of cursorial spider assemblages was examined along a gradient of four temperature successional communities. Species diversity (H), richness (S), and evenness (J) exhibited a dichotomy between herbaceous and woody communities rather than a progressive change with community age: all three parameters were higher in the two younger fields than in the two older woodlands, which is contrary to conventional successional theory. Species importance curves were steeper in the two woody communities. The breadth of the distribution of adult body lengths was greater in the two herbaceous communities. Indices of community similarity revealed neither a successional trend nor the vegetative dichotomy. We suggest the hypothesis that habitat structure is a more important determinant of cursorial spider diversity than successional age per se, and that the switch in dominance from herbaceous to woody vegetation is the critical change. We further suggest that competition for prey is more important to cursorial spiders in early successional (herbaceous) communities, because of a switch in the limiting resource from prey in these communities to the amount of accumulating litter (a spatial resource) in older woody stands. This may explain the greater variation in adult body size of these generalist predators in the two younger communities.     

Diet-Dependent Survival, Development and Fecundity of the Spider Atypena formosana (Oi) (Araneae: Linyphiidae)Implications for Biological Control in Rice

The development time of Atypena formosana was assessed on four different diets: (1) brown planthopper (BPH), Nilaparvata lugens ; (2) green leafhopper (GLH), Nephotettix virescens ; (3) Collembola (Entomobryidae); and (4) a mixed diet of Collembola, hoppers and Drosophila melanogaster . A starvation treatment served as a control. We tested the hypothesis that because of differences in food quality, the diets would affect survival and development differently. Survival differed significantly among the diets. No spiders survived until adult on the diet of GLH, only a single individual survived on the diet of BPH, whereas 70% survived until adult on the Collembola diet and all survived on the mixed diet. Development times were significantly different among the diets tested. Spiders developed fastest on the mixed diet and the Collembola diet. The biomass of prey consumed within 48 h and the total biomass of prey consumed to complete an instar were highest on the mixed diet. Finally the mixed diet resulted in higher fecundity than the Collembola diet. It is concluded that alternative prey are an absolute necessity, as BPH and GLH diets alone would not be able to increase the number of predators, regardless of the number of hoppers present.