Distinct feeding strategies of generalist and specialist spiders

1. Feeding behaviour of generalist and specialist predators is determined by a variety of trophic adaptations. Specialised prey‐capture adaptations allow specialists to catch relatively large prey on a regular basis. As a result, specialists might be adapted to exploit each item of prey more thoroughly than do generalists. 2. It was expected that obligatory specialist cursorial spiders would feed less frequently than generalists but for a longer time and, thus, that their foraging pause would be longer. First, the feeding frequencies of three generalist spider species (Cybaeodamus taim, Harpactea hombergi, Hersiliola sternbergsi) were compared with those three phylogenetically related specialist species: myrmecophagous Zodarion rubidum, and araneophagous Nops aff. variabilis and Palpimanus orientalis. 3. Generalists captured more prey, exploited each item of prey for a significantly shorter time, and had a shorter foraging pause than was the case for specialists. Generalists also gained significantly less relative amount of prey mass than did specialists. 4. Second, the study compared the prey DNA degradation rate in the gut of generalists and specialists by means of PCR. The degradation rate was not significantly different between specialists and generalists: the detectability half‐life was estimated to exist for 14.3 days after feeding. 5. This study shows that the feeding strategies of cursorial generalist and obligatory specialist spiders are different. Obligatory specialists have evolved a feeding strategy that is based on thorough exploitation of a few large prey, whereas generalists have evolved a strategy that is based on short exploitation of multiple small items of prey.     

Host specificity in phytophagous insects: selection pressure from generalist predators

The narrow host range of insect herbivores is noted, and some of the reasons why specificity has evolved are reviewed. Works indicating the need for new approaches are pointed out including the possibility that generalist predators provide a suitable pressure. Experiments to test the hypothesis that generalists are more vulnerable than specialists to predators are described. They involved a vespid wasp and over thirty species pairs of caterpillars, matched for size and density. Overall, generalists were taken more readily than specialists: some but not all reasons could be detected. The results are discussed in ecological and evolutionary terms.     

Response of pest control by generalist predators to local‐scale plant diversity: a meta‐analysis

Disentangling the effects of plant diversity on the control of herbivores is important for understanding agricultural sustainability. Recent studies have investigated the relationships between plant diversity and arthropod communities at the landscape scale, but few have done so at the local scale. We conducted a meta‐analysis of 32 papers containing 175 independent measures of the relationship between plant diversity and arthropod communities. We found that generalist predators had a strong positive response to plant diversity, that is, their abundance increased as plant diversity increased. Herbivores, in contrast, had an overall weak and negative response to plant diversity. However, specialist and generalist herbivores differed in their response to plant diversity, that is, the response was negative for specialists and not significant for generalists. While the effects of scale remain unclear, the response to plant diversity tended to increase for specialist herbivores, but decrease for generalist herbivores as the scale increased. There was no clear effect of scale on the response of generalist predators to plant diversity. Our results suggest that the response of herbivores to plant diversity at the local scale is a balance between habitat and trophic effects that vary according to arthropod specialization and habitat type. Synthesis and applications. Positive effects of plant diversity on generalist predators confirm that, at the local scale, plant diversification of agroecosystems is a credible and promising option for increasing pest regulation. Results from our meta‐analysis suggest that natural control in plant‐diversified systems is more likely to occur for specialist than for generalist herbivores. In terms of pest management, our results indicate that small‐scale plant diversification (via the planting of cover crops or intercrops and reduced weed management) is likely to increase the control of specialist herbivores by generalist predators.     

Elimination of plant toxins by herbivorous woodrats: revisiting an explanation for dietary specialization in mammalian herbivores

Constraints on rates of detoxification and elimination of plant toxins are thought to be responsible for limiting dietary specialization in mammalian herbivores. This hypothesis, known as the detoxification limitations hypothesis, suggests that most mammalian herbivores are generalists to avoid overdosing on toxins from a single plant species. The hypothesis also predicts that the few mammalian specialists that exist should have adaptations for rapid detoxification and elimination of plant secondary compounds. We took a pharmacological approach to test whether specialists eliminate toxins from the bloodstream faster than generalists. We compared elimination rate and total exposure of alpha-pinene in closely related dietary specialist and generalist woodrats, Neotoma stephensi and N. albigula, respectively. Animals were orally gavaged with alpha-pinene, a plant secondary compound present in the natural diets of both woodrat species. We collected venous blood at 3, 6, 10, 15, and 20 min post-ingestion of alpha-pinene. Blood was analyzed for alpha-pinene concentration using gas chromatography. We found that specialist and generalist woodrats did not differ in elimination rates of alpha-pinene. However, specialists had lower exposure levels of alpha-pinene than generalists due to lower initial delivery of alpha-pinene to the general circulation. The levels of alpha-pinene detected in the bloodstream of specialists were 4.7-5.3x lower over all time intervals than generalists. Thus, specialists encounter a functionally lower dose of toxin than generalists. We suggest that the lower exposure level of specialist woodrats may be due to mechanisms in the gut that decrease toxin absorption. Regardless of mechanism, lower exposure to plant toxins may allow specialists to forage on diets with high toxin concentrations thereby facilitating dietary specialization.     

Generalist caterpillar prey are more palatable than specialists for the generalist predator Iridomyrmex humilis

Summary Experiments are described which test the hypothesis that more host-specific species of caterpillars should be less aceptable to a generalist predator than polyphagous species. Caterpillars of all species were tested in paired choice tests with the Argentine ant, Iridomyrmex humilis. Experiments were replicated ten times, videotaped and later analyzed. Brightly colored specialist species (normally considered to be aposematic) were clearly the least palatable, while more cryptic specialists were also significantly less acceptable than generalists overall. Leaf-tying species were considered separately; all were highly palatable independent of host range. The results indicate that among caterpillars that do not construct leaf shelters, those with a wide post range are more acceptable than those with a narrow host range. This is consistent with the notion that generalist predators provide selection pressure favoring narrow host range in their herbivorous prey.     

Variation in the degree of specialization can maintain local diversity in model communities

We hypothesize that the continuum between generalist and specialist adaptations is an important general trade-off axis in the maintenance of local diversity, and we explore this idea with a simple model in which there are patch types to which species arrive as propagules and compete. Each patch type is defined by a competitive ranking of all species. A highly specialist species is the top competitor in one patch type but has a relatively low average ranking across different patch types, while a generalist species has a high average rank across patch types but is not the top competitor in any patch type. We use random dispersal and vary the fecundity of all species together to vary total propagule density and therefore recruitment limitation and density-dependent mortality. When fecundity is very high, each patch becomes occupied by its specialist species and generalists go extinct, so the number of species at equilibrium is equal to the number of patch types. If fecundity is very low, generalists dominate and specialists go extinct. There is a range of fecundity levels in which specialists, generalists, and intermediates coexist, and the number of species is substantially greater than the number of patch types. While coexistence of specialists and generalists has been considered a problem in evolutionary ecology, our results suggest to the contrary that this trade-off contributes to the maintenance of local diversity.     

Increased genetic differentiation in a specialist versus a generalist bee: implications for conservation

Oligolectic bees are specialists that collect pollen from one or a few closely related species of plants, while polylectic bees are generalists that collect pollen from both related and unrelated species of plants. Because of their more restricted range of floral hosts, it is expected that specialists persist in more isolated populations than do generalists. We present data on the population structure of two closely related bee species sampled from a super abundant floral host in the southern Atacama Desert. Pairwise comparisons of population subdivision over identical distances revealed that the specialist bee had significantly more differentiated populations in comparison to the generalist. Further, populations of the specialist had significantly less genetic variation, measured as observed and expected heterozgyosity, than those of the generalist. Our data support the hypothesis of decreased gene flow among populations of the specialist bee even at equivalent geographic distances. The resulting reductions in effective population size for specialists make them particularly prone to extinction due to both demographic and genetic reasons. Our findings have important implications for the conservation of bees and other specialist insects. Deceased     

Microbial generalist or specialist: Intraspecific variation and dormancy potential matter

Microbial generalists and specialists coexist in the soil environment while having distinctive impacts on microbial community dynamics. In microbial ecology, the underlying mechanisms as to why a species is a generalist or a specialist remain ambiguous. Herein, we collected soils across a national scale and identified bacterial generalists and specialists according to niche breadth at the species level (OTU level), and the single-nucleotide differences in each species were measured to investigate intraspecific variation (at zero-radius OTU level). Compared with that of the specialists, the intraspecific variation of the generalists was much higher, which ensured their wider niche breadth and lower variability. The higher asynchrony and different niche preferences of conspecific individuals and the higher dormancy potential within the generalists further contributed to their stability in varying environments. Besides, generalists were less controlled by environmental filtering, which was indicated by the stronger signature of stochastic processes in their assembly, and had higher diversification and transition rates that allowed them to adapt to environmental changes to a greater extent than specialists. Overall, this study provides a new comprehensive understanding of the rules of assembly and the evolutionary roles of bacterial generalists and specialists. It also highlights the importance of intraspecific variation and the dormancy potential in the stability of species.     

Specialized avian Haemosporida trade reduced host breadth for increased prevalence

Parasite specialization on one or a few host species leads to a reduction in the total number of available host individuals, which may decrease transmission. However, specialists are thought to be able to compensate by increased prevalence in the host population and increased success in each individual host. Here, we use variation in host breadth among a community of avian Haemosporida to investigate consequences of generalist and specialist strategies on prevalence across hosts. We show that specialist parasites are more prevalent than generalist parasites in host populations that are shared between them. Moreover, the total number of infections of generalist and specialist parasites within the study area did not vary significantly with host breadth. This suggests that specialists can infect a similar number of host individuals as generalists, thus compensating for a reduction in host availability by achieving higher prevalence in a single host species. Specialist parasites also tended to infect older hosts, whereas infections by generalists were biased towards younger hosts. We suggest that this reflects different abilities of generalists and specialists to persist in hosts following infection. Higher abundance and increased persistence in hosts suggest that specialists are more effective parasites than generalists, supporting the existence of a trade‐off between host breadth and average host use among these parasites.     

Specialist and generalist herbivores exert opposing selection on a chemical defense

* Plant defense traits often show high levels of genetic variation, despite clear impacts on plant fitness. This variation may be partly maintained by trade-offs in the defense against multiple herbivore species, for example between generalists and coevolved specialists. Despite a long-standing discussion in the literature on the subject, no study to date has specifically manipulated specialist and generalist herbivores independently of one another to determine whether the two guilds exert opposing selection pressures on specific defensive traits. * In two separate experiments, the dominant specialist and generalist herbivores of Brassica nigra were independently manipulated to test whether the composition of the herbivore community altered the direction of selection on a major defensive trait of the plant, sinigrin concentration. * It was found that generalist damage was negatively correlated but specialist loads were positively correlated with increasing sinigrin concentrations; and sinigrin concentration was favored when specialists were removed, disfavored (past an intermediate point) when generalists were removed and selectively neutral when plants faced both generalists and specialists.     

Enemy release but no evolutionary loss of defence in a plant invasion: an inter-continental reciprocal transplant experiment

Plant chemical defenses and escape from natural enemies have been postulated to select for dietary specialization in herbivorous insects. In field and laboratory bioassays, we evaluated the effectiveness of intact and chemically modified larval shield defenses of the generalist Chelymorpha alternans and the specialists Acromis sparsa and Stolas plagiata (Chrysomelidae: Cassidinae) against three natural predators, using larvae reared on two morning glory (Convolvulaceae) species. We assessed whether: (1) specialists were better defended than generalists when both were fed and assayed on the same plant; (2) larval shield defenses were chemical, physical, or both; and (3) specialists exploit chemistry better than generalists. Live specialist larvae survived at higher rates than did generalists in predator bioassays with the bug Montina nigripes (Reduviidae), but there were no differences among groups against two species of Azteca ants (Hymenoptera: Dolichoderinae). Solvent leaching by H₂O or MeOH significantly reduced shield efficacy for all species compared to larvae with intact shields. In contrast, freshly killed specialist larvae exhibited significantly lower capture rates and frequencies than the generalists. Although solvent leaching significantly reduced overall shield efficacy for freshly killed larvae of all species, the pattern of leaching effects differed between specialists and generalists, with H₂O-leaching having a greater impact on the specialists. The overall vulnerability of the generalists appears due to lower chemical protection, which is ameliorated by increased escape behaviors, suggesting a selective trade-off between these defensive components. These experiments indicate that shield defenses are essential for larval survival and that specialists are superior at exploiting plant compounds residing in the aqueous fraction. Our results support the hypothesis that diet-specialized herbivorous insects have more effective defenses than generalists when both feed on the same plant due to the differential ability to exploit defensive precursors obtained from the host. The evolution of dietary specialization may therefore confer the advantage of enhanced enemy-free space.     

Spider cocoons and their suspension systems as barriers to generalist and specialist predators

Summary This study tests hypotheses regarding spider cocoons and their suspension systems as barriers to generalist and specialist predators. Evidence presented here suggests that the suspension systems ofMecynogea lemniscata andArgiope aurantia cocoons are effective barriers against small generalists such as ants, but fail to stop large generalists such as birds. Cocoon covers were found to be generally ineffective against generalist predators. Various component layers of these cocoons are shown, however, to be an effective barrier against the attack modes of specific predator guilds. Cocoon covers function primarily as barriers to specialists that use active “burrowing” larvae to gain entrance into the cocoon, while the flocculent silk layer is shown to be an effective barrier against specialists which use a long ovipositor to attack cocoons. These findings support suggestions that the primary role of the cocoon is to provide protection from predators and parasites. These results also support a close evolutionary relationship between cocoon architecture and specialized predators.     

The tri-trophic interactions hypothesis: interactive effects of host plant quality, diet breadth and natural enemies on herbivores

Several influential hypotheses in plant-herbivore and herbivore-predator interactions consider the interactive effects of plant quality, herbivore diet breadth, and predation on herbivore performance. Yet individually and collectively, these hypotheses fail to address the simultaneous influence of all three factors. Here we review existing hypotheses, and propose the tri-trophic interactions (TTI) hypothesis to consolidate and integrate their predictions. The TTI hypothesis predicts that dietary specialist herbivores (as compared to generalists) should escape predators and be competitively dominant due to faster growth rates, and that such differences should be greater on low quality (as compared to high quality) host plants. To provide a preliminary test of these predictions, we conducted an empirical study comparing the effects of plant (Baccharis salicifolia) quality and predators between a specialist (Uroleucon macolai) and a generalist (Aphis gossypii) aphid herbivore. Consistent with predictions, these three factors interactively determine herbivore performance in ways not addressed by existing hypotheses. Compared to the specialist, the generalist was less fecund, competitively inferior, and more sensitive to low plant quality. Correspondingly, predator effects were contingent upon plant quality only for the generalist. Contrary to predictions, predator effects were weaker for the generalist and on low-quality plants, likely due to density-dependent benefits provided to the generalist by mutualist ants. Because the TTI hypothesis predicts the superior performance of specialists, mutualist ants may be critical to A. gossypii persistence under competition from U. macolai. In summary, the integrative nature of the TTI hypothesis offers novel insight into the determinants of plant-herbivore and herbivore-predator interactions and the coexistence of specialist and generalist herbivores.     

Suitability of woodlice prey for generalist and specialist spider predators: a comparative study

1. Predators select a prey according to its energetic and nutritional composition. Generalist predators avoid, whereas specialists often specialise on well‐defended prey. The aim of this study was to find the suitability of woodlice prey for generalist and specialist predators by comparing their handling efficiency. 2. Laboratory experiments were performed in which specialist and generalist predators were reared on monotypic diets comprising one or other of two woodlice species that differ in their defensive strategies: rollers (Armadillidium) and clingers (Porcellio). A control group was reared on a mixture of arthropods (excluding woodlice). Three spider predators were used that differ in their adaptations to deal with woodlice prey: a woodlice specialist, Dysdera crocata; an oligophagous generalist, Pholcus phalangioides, that also captures woodlice; and a euryphagous generalist, Tegenaria domestica, that does not feed on woodlice. The frequency of capture was recorded and various fitness parameters were measured, namely survival, growth rate, and ontogenetic development. 3. It was found that the specialist, D. crocata, performed best on the Porcellio diet, and similarly well on Armadillidium and mixed diets. The two generalists, P. phalangioides and T. domestica, had poor performance on both woodlice diets but performed well on the mixed diet. 4. The results show that woodlice are unsuitable prey for both oligophagous and euryphagous generalist predators.     

Coexistence of Habitat Specialists and Generalists in Metapopulation Models of Multiple-Habitat Landscapes

In coarse-grained environments specialists are generally predicted to dominate. Empirically, however, coexistence with generalists is often observed. We present a simple, but previously unrecognized, mechanism for coexistence of a habitat generalist and a number of habitat specialist species. In our model all species have a metapopulation structure in a landscape consisting of patches of different habitat types, governed by local extinction and colonization. Each specialist is limited to its specific type of habitat. The generalist can use more types of habitat, has a lower local competitive ability but can exploit patches left open by the specialists. Our modeling shows that coexistence is easily possible. The mechanism amounts to a colonization/competition trade-off at the landscape level, where the colonization advantage of the inferior competitor does not arise from a higher colonization rate but from its ability to use more types of habitat. Habitat availability has to be intermediate: when there are few patches of each habitat, only the generalist is able to maintain itself and when there are many patches, high propagule pressure of the specialists excludes the generalist. Habitat selection or temporal variations in relative habitat quality are not necessary for coexistence. Increased niche-width, colonization rate or local competitive ability of the generalist enhances its performance compared to the specialists. Various types of habitat degradation favour generalism. When able to use a broad range of habitats, generalists can generate so much propagule pressure that only a low level of local competitive ability is needed to globally exclude the specialists. Hence, in a reversal of the original problem, the question is why there are so many specialist metapopulations?     

Does habitat specialization shape the evolutionary potential of wild bird populations?

Because specialist species evolved in more temporally and spatially homogeneous environments than generalist species, they are supposed to experience less fluctuating selection. For this reason, we expect specialists to show lower overall genetic variation as compared to generalists. We also expect populations from specialist species to be smaller and more fragmented, with lower neutral genetic diversity. We tested these hypotheses by investigating patterns of genetic diversity along a habitat specialization gradient in wild birds, based on estimates of heritability, coefficients of variation of additive genetic variance, and heterozygosity available in the literature. We found no significant effect of habitat specialization on any of the quantitative genetic estimators but generalists had higher heterozygosity. This effect was mainly a consequence of the larger population size of generalists. Our results suggest that evolutionary potential does not differ at the population level between generalist and specialist species, but the trend observed in heterozygosity levels and population sizes may explain their difference in susceptibility to extinction.     

Eating their way to the top? Mechanisms underlying the success of invasive insect generalist predators

Insect generalist predators have been introduced outside of their native range intentionally to improve biological control, or accidentally during commerce, and can subsequently become invasive. Invasive insect generalist predators (IIGP) have widespread impacts on invaded communities because they consume both herbivores and other predators. Also, they often reach higher densities than and displace similar native species. Reflecting the complexity of their ecological roles, a wide variety of mechanisms might contribute to invasive success by IIGP. These species often drive resources to lower levels than do natives, leading to intense resource competition and sometimes competitive exclusion of other predators. The broader range of resources used by many IIGP can heighten their competitive advantage, particularly when IIGP exploit modified habitats. In either case, IIGP improve herbivore suppression by depressing prey densities below pre-invasion levels. Coexistence among native and invasive generalists is fostered when species differ substantially in their niche requirements. In this case, a larger proportion of the total prey population is subject to attack post- than pre-invasion, which strengthens prey suppression. On the other hand, some IIGP feed heavily on other predators, at times leading to a weakening of prey suppression. Future research should continue to explore the roles of competition and niche partitioning on larger spatial scales, and in both the native and invasive ranges of IIGP. Additionally, combining data from empirical studies with theory might be an effective way to predict the spread and community impacts of IIGP invasions.     

Cannibalism and Intraguild Predation Among Phytoseiid Mites: Are Aggressiveness and Prey Preference Related to Diet Specialization?

We tested whether specialist and generalist phytoseiid mites differ in aggressiveness and prey choice in cannibalism and intraguild predation. Specialists tested were Galendromus occidentalis, Neoseiulus longispinosus, Phytoseiulus persimilis, and P. macropilis; tested were Amblyseius andersoni, Euseius finlandicus, E. hibisci, Kampimodromus aberrans, Neoseiulus barkeri, N. californicus, N. cucumeris, N. fallacis, and Typhlodromus pyri. Aggressiveness of cannibalistic females against larvae was not related to diet specialization except that highly aggressive species were exclusively generalists. Seldom to moderately cannibalizing species occurred in both generalist and specialist phytoseiids. In contrast to aggressiveness in cannibalism, generalists and specialists differed in aggressiveness in intraguild predation. Adult females of specialists were only slightly aggressive against heterospecific larvae, whereas adult females of all generalists except T. pyri were highly aggressive. Adult females of generalists were able to discriminate between con- and heterospecific larvae and preferentially consumed the latter when given a choice. Adult females of specialists except G. occidentalis showed no preference when given a choice between con- and heterospecific larvae. We conclude that aggressiveness in intraguild predation, species recognition and subsequent preferential consumption of heterospecifics when given a choice is common in generalist but not specialist phytoseiids. We discuss the evolutionary pathways that might have led to the difference between specialists and generalists in species discrimination.     

Numerical and dietary responses of a predator community in a temperate zone of Europe

The generalist predation hypothesis predicts that the functional responses of generalist predator species should be quicker than those of specialist predators and have a regulating effect on vole populations. New interpretations of their role in temperate ecosystems have, however, reactivated a debate suggesting generalist predators may have a destabilizing effect under certain conditions (e.g. landscape homogeneity, low prey diversity, temporary dominance of 1 prey species associated with a high degree of dietary specialization). We studied a rich predator community dominated by generalist carnivores ( Martes spp., Vulpes vulpes, Felis catus ) over a 6 yr period in farmland and woodland in France. The most frequent prey were small rodents (mostly Microtus arvalis , a grassland species, and Apodemus spp., a woodland species). Alternative prey were diverse and dominated by lagomorphs ( Oryctolagus cuniculus, Lepus europeus ). We detected a numerical response among specialist carnivores but not among generalist predators. The dietary responses of generalist predators were fairly complex and most often dependent on variation in density of at least 1 prey species. These results support the generalist predation hypothesis. We document a switch to alternative prey, an increase of diet diversity, and a decrease of diet overlap between small and medium-sized generalists during the low density phase of M. arvalis . In this ecosystem, the high density phases of small mammal species are synchronous and cause a temporary specializing of several generalist predator species. This rapid functional response may indicate the predominant role of generalists in low amplitude population cycles of voles observed in some temperate areas.     

Landscape structure and agricultural intensification are weak predictors of host range and parasitism rate of cereal aphids

Proportions of specialist and generalist primary parasitoids have been described by the resource breadth and the trade-off hypothesis. These alternative hypotheses predict either decreased or increased, respectively, parasitism rate of shared aphid species by specialist parasitoids. We tested both hypotheses and the confounding effects of landscape structure and agricultural intensification (AI) using extensive samplings of aphids and their parasitoids in Polish agricultural landscapes. Abundances, species composition of aphids, primary parasitoids, and parasitism rate of aphids by specialists and generalist parasitoids were analysed. Contrary to our expectations we found equally decreased parasitism rates by both types of primary parasitoids at higher aphid densities and thus proportion of specialists to generalists did not change with increasing host density. In line with the resource breadth hypothesis, specialist parasitoids had always lower abundances and parasitism rates than generalist parasitoids. Landscape diversity and agricultural intensification did not influence the host-parasitoid population dynamics. We speculate that these contrasting results could be caused by the additional density effects of secondary parasitoids. We conclude that simplistic two-trophic-level population models are not able to fully describe the complex dynamics of trophic networks. We also argue that agricultural intensification has lower effects on abundance and effectiveness of parasitoids than predicted by respective predator–prey models and empirical studies performed in controlled and artificial conditions.