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1.
Terrestrial predators on marine shores benefit from the inflow of organisms and matter from the marine ecosystem, often causing very high predator densities and indirectly affecting the abundance of other prey species on shores. This indirect effect may be particularly strong if predators shift diets between seasons. We therefore quantified the seasonal variation in diet of two wolf spider species that dominate the shoreline predator community, using molecular gut content analyses with general primers to detect the full prey range. Across the season, spider diets changed, with predominantly terrestrial prey from May until July and predominantly marine prey (mainly chironomids) from August until October. This pattern coincided with a change in the spider age and size structure, and prey abundance data and resource selection analyses suggest that the higher consumption of chironomids during autumn is due to an ontogenetic diet shift rather than to variation in prey abundance. The analyses suggested that small dipterans with a weak flight capacity, such as Chironomidae, Sphaeroceridae, Scatopsidae and Ephydridae, were overrepresented in the gut of small juvenile spiders during autumn, whereas larger, more robust prey, such as Lepidoptera, Anthomyidae and Dolichopodidae, were overrepresented in the diet of adult spiders during spring. The effect of the inflow may be that the survival and growth of juvenile spiders is higher in areas with high chironomid abundances, leading to higher densities of adult spiders and higher predation rates on the terrestrial prey next spring.  相似文献   

2.
Aim We compiled data on prey utilization of spiders at a global scale to better understand the relationship between current climate or net primary production (NPP) and diet breadth, evenness and composition in spiders. We test whether the productivity and the diversity–climatic‐stability (DCS) hypotheses focusing on diversity patterns may also explain global patterns in prey utilization by web‐building and cursorial spiders. Location A global dataset of 95 data points from semi‐natural and natural terrestrial habitats spanning 41.3° S to 56.1° N. Methods We collected data on spider prey (29 groups, mostly order‐level invertebrate taxa) through extensive literature research to identify the relationship between climatic conditions and NPP and spider diets based on 66 studies of prey composition in 82 spider species. Results The number of prey groups in spider diets was positively related to NPP, after accounting for differences in sampling effort in the original studies. In general, diet breadth was significantly higher for spider species in tropical environments. Prey individuals in spider diets were more evenly distributed among different prey groups in warmer environments with lower fluctuations in precipitation. Collembola and other spiders were more common prey for spiders with a cursorial hunting mode. Myriapoda and Collembola were more common prey in cooler climates with more stable precipitation, whereas Isoptera, Lepidoptera, Psocoptera and Coleoptera showed the opposite pattern. Main conclusions The positive relationship between diet breadth and NPP and the negative relationship between prey evenness and seasonality in precipitation support the productivity and the DCS hypotheses, respectively. This effect on global patterns of invertebrate predator–prey interactions suggests that trophic interactions between spiders and their prey are sensitive to climatic conditions. Climatic conditions may not only affect spider community composition, but also considerably alter the functional role of these abundant invertebrate predators in terrestrial ecosystems.  相似文献   

3.
Spiders are dominant terrestrial predators that consume a large variety of prey and engage in intraguild predation. Although the feeding habits of certain species are well known, the trophic structure of spider assemblages still needs to be investigated. Stable isotope analysis enables characterisation of trophic relationships between organisms because it tracks the energy flow in food webs and indicates the average number of trophic transfers between a given species and the base of the web, thus being a useful tool to estimate the magnitude of intraguild predation in food webs. Using this technique, we studied the trophic groups of spiders and their links within the arthropod food web of a Mediterranean organic citrus grove. We assessed the trophic positions of the 25 most common spider species relative to other arthropod predators and potential prey in the four seasons of the year, both in the canopy and on the ground. The analyses showed great seasonal variation in the isotopic signatures of some arthropod species, as well as the existence of various trophic groups and a wide range of trophic levels among spiders, even in species belonging to the same family. Differences in δ15N between spiders and the most abundant prey in the grove usually spanned two trophic levels or more. Our findings provide field evidence of widespread intraguild predation in the food web and caution against using spider families or guilds instead of individual species when studying spider trophic interactions.  相似文献   

4.
Specialist true predators are expected to exhibit higher capture efficiencies for the capture of larger and dangerous prey than generalist predators due to their possession of specialized morphological and behavioral adaptations. We used an araneophagous spider (Lampona murina) and a generalist spider (Drassodes lapidosus) as phylogenetically related model species and investigated their realized and fundamental trophic niches and their efficacy with respect to prey capture and prey handling. The trophic niche of both species confirmed that Lampona had a narrow trophic niche with a predominance of spider prey (including conspecifics), while the niche of Drassodes was wide, without any preference. DNA analysis of the gut contents of Lampona spiders collected in the field revealed that spiders form a significant part of its natural diet. Lampona captured significantly larger prey than itself and the prey captured by Drassodes. As concerns hunting strategy, Lampona grasped the prey with two pairs of legs possessing scopulae, whereas Drassodes immobilized prey with silk. Lampona possess forelegs equipped with scopulae and a thicker cuticle similar to other nonrelated araneophagous spiders. Lampona fed for a longer time and extracted more nutrients than Drassodes. We show that specialized behavioral and morphological adaptations altogether increase the hunting efficiency of specialists when compared to generalists.  相似文献   

5.
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.  相似文献   

6.
Jeff Scott Wesner 《Oikos》2012,121(1):53-60
Food webs in different ecosystems are often connected through spatial resource subsidies. As a result, biodiversity effects in one ecosystem may cascade to adjacent ecosystems. I tested the hypothesis that aquatic predator diversity effects cascade to terrestrial food webs by altering a prey subsidy (biomass and trophic structure of emerging aquatic insects) entering terrestrial food webs, in turn altering the distribution of a terrestrial consumer (spider) that feeds on emerging aquatic insects. Fish presence, but not diversity, altered the trophic structure of emerging aquatic insects by strongly reducing the biomass of emerging predators (dragonflies) relative to non‐feeding taxa (chironomid midges). Fish diversity reduced emerging insect biomass through enhanced effects on the most common prey taxa: predatory dragonflies Pantala flavescens and non‐feeding chironomids. Terrestrial spiders (Tetragnathidae) primarily captured emerging chironomids, which were reduced in the high richness (3 spp.) treatment relative to the 1 and 2 species treatments. As a result, terrestrial spider abundance was lower above pools with high fish richness (3 species) than pools with 1 and 2 species. Synergistic predation effects were mostly limited to the high richness treatment, in which fish occupied each level of vertical microhabitat in the water‐column (benthic, middle, surface). This study demonstrates that predator diversity effects are not limited to the habitat of the predator, but can propagate to adjacent ecosystems, and demonstrates the utility of using simple predator functional traits (foraging domain) to more accurately predict the direction of predator diversity effects.  相似文献   

7.
Many ecosystems are linked to their adjacent ecosystems by movements of organisms. For instance, aquatic and terrestrial ecosystems are linked via emerging aquatic insects that serve as prey for terrestrial consumers. However, the role of these organisms in returning recycled carbon to the ecosystem from which it originated is not well known. This is due to the fact that values of carbon isotope signatures from terrestrial leaves and aquatic resources are usually similar and hence results of isotope mixing models need to be considered with caution. We overcame this problem by adding isotopically distinct terrestrial particulate organic carbon (tPOC) as a tracer to the experimental sides of two lakes that were divided in two equal halves with plastic curtains. We focused on aquatic insect larvae (Chironomidae) that fed on maize Zea mays leaves experimentally added to the lakes, and subsequently became prey for terrestrial predators (spiders) after emergence. The carbon isotope values of Chironomidae and spiders were significantly elevated in the lake treatment sides as compared to reference sides, whereas the values of all autochthonous resources were not affected by maize additions. Estimates from stable isotope mixing models indicated a low but demonstrable contribution of maize leaves to the diet of Chironomidae. Overlap between the isotope values of alder leaves, the major natural tPOC source, and autochthonous resources prevented a reliable quantification of allochthony of Chironomidae. However, we qualitatively demonstrated the flow of terrestrial particulate organic carbon to lakes, as leaf fall, and back to terrestrial surroundings via emerging insects. This ‘boomerang’ carbon flux between land and lakes blurs the distinction between autochthonous and allochthonous carbon sources.  相似文献   

8.
Predators appear to be less frequently specialised (i.e. adapted to restricted diet) on their prey than herbivores, parasites or parasitoids. Here, we critically evaluate contemporary evolutionary hypotheses that might be used to explain the evolution of specialised foraging in predators. We propose a unifying concept within which we define four types of trophic categories using ecological (diet breadth) and evolutionary (degree of adaptations) contexts. We use data on spiders (Araneae), the most diversified order of terrestrial predators, to assess applicability of frameworks and evolutionary concepts related to trophic specialisation. The majority of spider species are euryphagous but a few have a restricted prey range, i.e. they are stenophagous. We provide a detailed overview of specialisation on different prey types, namely spiders, crustaceans, moths, dipterans, ants, and termites. We also review the available evidence for trophic adaptations, classified into four categories: behavioural, morphological, venomic and metabolic. Finally, we discuss the ecological and evolutionary implications of trophic specialisation and propose avenues for future research.  相似文献   

9.
We examined the relationship between body size of the riparian spider Nephila clavata and the contribution of allochthonous (aquatic insects) and autochthonous (terrestrial insects) sources to its diet using stable isotope analysis. During the study period from July to September, the body size of the females increased remarkably (about 60-fold) but that of males remained small. The biomass of both aquatic and terrestrial insects trapped on the spider webs increased with spider size, with the biomass of the former ranging between 30 and 70% of that of the terrestrial insects. The average relative contribution of aquatic insects to the diet of the spiders, calculated from δ13C values, was 40–50% in spiders in the early juvenile and juvenile stages, 35% in adult males and 4% in adult females. There was a significant negative relationship between the relative contribution of aquatic insects and body size of the female spiders. We conclude that aquatic insects might be an important seasonal dietary subsidy for small spiders and that these allochthonous subsidies may facilitate the growth of riparian spiders, which may in turn enable the spiders to feed on larger prey.  相似文献   

10.
11.
Several hypotheses have been put forward to explain the evolution of prey specificity (stenophagy). Yet little light has so far been shed on the process of evolution of stenophagy in carnivorous predators. We performed a detailed analysis of a variety of trophic adaptations in one species. Our aim was to determine whether a specific form of stenophagy, myrmecophagy, has evolved from euryphagy via parallel changes in several traits from pre-existing characters. For that purpose, we studied the trophic niche and morphological, behavioural, venomic and physiological adaptations in a euryphagous spider, Selamia reticulata. It is a species that is branching off earlier in phylogeny than stenophagous ant-eating spiders of the genus Zodarion (both Zodariidae). The natural diet was wide and included ants. Laboratory feeding trials revealed versatile prey capture strategies that are effective on ants and other prey types. The performance of spiders on two different diets – ants only and mixed insects – failed to reveal differences in most fitness components (survival and developmental rate). However, the weight increase was significantly higher in spiders on the mixed diet. As a result, females on a mixed diet had higher fecundity and oviposited earlier. No differences were found in incubation period, hatching success or spiderling size. S. reticulata possesses a more diverse venom composition than Zodarion. Its venom is more effective for the immobilisation of beetle larvae than of ants. Comparative analysis of morphological traits related to myrmecophagy in the family Zodariidae revealed that their apomorphic states appeared gradually along the phylogeny to derived prey-specialised genera. Our results suggest that myrmecophagy has evolved gradually from the ancestral euryphagous strategy by integrating a series of trophic traits.  相似文献   

12.
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.  相似文献   

13.
A forest-stream trophic link was examined by stable carbon isotope analyses which evaluated the relationship of aquatic insects emerging from a stream to the diets of web-building spiders. Spiders, aquatic and terrestrial prey, and basal resources of forest and stream food webs were collected in a deciduous forest along a Japanese headwater stream during May and July 2001. The 13C analyses suggested that riparian tetragnathid spiders relied on aquatic insects and that the monthly variation of such dependence is partly associated with the seasonal dynamics of aquatic insect abundance in the riparian forest. Similarly, linyphiid spiders in the riparian forest exhibited 13C values similar to aquatic prey in May. However, their 13C values were close to terrestrial prey in both riparian and upland (150m away from the stream) forests during June to July, suggesting the seasonal incorporation of stream-derived carbon into their tissue. In contrast, araneid spiders relied on terrestrial prey in both riparian and upland forests throughout the study period. These isotopic results were consistent with a previous study that reported seasonal variation in the aquatic prey contribution to total web contents for each spider group in this forest, implying that spiders assimilate trapped prey and that aquatic insect flux indeed contributes to the energetics of riparian tetragnathid and linyphiid spiders.  相似文献   

14.
Although some primary consumers such as chironomid larvae are known to exploit methane‐derived carbon via microbial consortia within aquatic food webs, few studies have traced the onward transfer of such carbon to their predators. The ruffe Gymnocephalus cernuus is a widespread benthivorous fish which feeds predominantly on chironomid larvae and is well adapted for foraging at lower depths than other percids. Therefore, any transfer of methanogenic carbon to higher trophic levels might be particularly evident in ruffe. We sampled ruffe and chironomid larvae from the littoral, sub‐littoral and profundal areas of Jyväsjärvi, Finland, a lake which has previously been shown to contain chironomid larvae exhibiting the very low stable carbon isotope ratios indicative of methane exploitation. A combination of fish gut content examination and stable isotope analysis was used to determine trophic linkages between fish and their putative prey. Irrespective of the depth from which the ruffe were caught, their diet was dominated by chironomids and pupae although the proportions of taxa changed. Zooplankton made a negligible contribution to ruffe diet. A progressive decrease in δ13C and δ15N values with increasing water column depth was observed for both chironomid larvae and ruffe, but not for other species of benthivorous fish. Furthermore, ruffe feeding at greater depths were significantly larger than those feeding in the littoral, suggesting an ontogenetic shift in habitat use, rather than diet, as chironomids remained the predominant prey item. The outputs from isotope mixing models suggested that the incorporation of methane‐derived carbon to larval chironomid biomass through feeding on methanotrophic bacteria increased at greater depth, varying from 0% in the littoral to 28% in the profundal. Using these outputs and the proportions of littoral, sub‐littoral or profundal chironomids contributing to ruffe biomass, we estimated that 17% of ruffe biomass in this lake was ultimately derived from chemoautotrophic sources. Methanogenic carbon thus supports considerable production of higher trophic levels in lakes.  相似文献   

15.
Studies on resource sharing and partitioning generally consider species that occur in the same habitat. However, subsidies between linked habitats, such as streams and riparian zones, create potential for competition between populations which never directly interact. Evidence suggests that the abundance of riparian consumers declines after fish invasion and a subsequent increase in resource sharing of emerging insects. However, diet overlap has not been investigated. Here, we examine the trophic niche of native fish, invasive fish, and native spiders in South Africa using stable isotope analysis. We compared spider abundance and diet at upstream fishless and downstream fish sites and quantified niche overlap with invasive and native fish. Spider abundance was consistently higher at upstream fishless sites compared with paired downstream fish sites, suggesting that the fish reduced aquatic resource availability to riparian consumers. Spiders incorporated more aquatic than terrestrial insects in their diet, with aquatic insects accounting for 45–90% of spider mass. In three of four invaded trout rivers, we found that the average proportion of aquatic resources in web‐building spider diet was higher at fishless sites compared to fish sites. The probability of web‐building and ground spiders overlapping into the trophic niche of invasive brown and rainbow trout was as high as 26 and 51%, respectively. In contrast, the probability of spiders overlapping into the trophic niche of native fish was always less than 5%. Our results suggest that spiders share resources with invasive fish. In contrast, spiders had a low probability of trophic overlap with native fish indicating that the traits of invaders may be important in determining their influence on ecosystem subsidies. We have added to the growing body of evidence that invaders can have cross‐ecosystem impacts and demonstrated that this can be due to niche overlap.  相似文献   

16.
Numerous studies have found that predators can suppress prey densities and thereby impact important ecosystem processes such as plant productivity and decomposition. However, prey suppression by spiders can be highly variable. Unlike predators that feed on prey within a single energy channel, spiders often consume prey from asynchronous energy channels, such as grazing (live plant) and epigeic (soil surface) channels. Spiders undergo few life cycle changes and thus appear to be ideally suited to link energy channels, but ontogenetic diet shifts in spiders have received little attention. For example, spider use of different food channels may be highly specialized in different life stages and thus a species may be a multichannel omnivore only when we consider all life stages. Using stable isotopes, we investigated whether wolf spider (Pardosa littoralis, henceforth Pardosa) prey consumption is driven by changes in spider size. Small spiders obtained > 80% of their prey from the epigeic channel, whereas larger spiders used grazing and epigeic prey almost equally. Changes in prey consumption were not driven by changes in prey density, but by changes in prey use by different spider size classes. Thus, because the population size structure of Pardosa changes dramatically over the growing season, changes in spider size may have important implications for the strength of trophic cascades. Our research demonstrates that life history can be an important component of predator diet, which may in turn affect community- and ecosystem-level processes.  相似文献   

17.
Spiders are the most common and predominant predators in terrestrial ecosystems. The predatory behavior of spiders affects the energy flow across the food web within an ecosystem. Traditiaonal methods for analyzing spider diets such as field observation, anatomy and faeces analysis are not suitable for spider experiments due to spiders’ special dietary behavior. The molecular method based on the specific primers of prey DNA seems to be inefficient either in spite of its wide application in diet analysis. As the next-generation sequencing (NGS) technology becomes prevalent in many different areas, several cases of the NGS-based analysis of mammal diets have been published. This study analyzed the diet differences of Pardosa pseudoannulata (Araneae: Lycosidae) in four habitats (a wetland, a tea plantation, an alpine meadow and a paddy field) by using the NGS technology, combined with the DNA barcode method. The results suggested that the Pardosa pseudoannulata feed on a broad range of prey, and 7 orders and 24 families of insects were detected in the four investigated habitats. Moreover, it is found that the diet diversity of Pardosa pseudoannulata is greatly influenced by their living environments and seasons. In a nutshell, this study established an NGS-based methodology for spider diets analysis, and the results provided some basic materials to inform the protection and utilization of the Pardosa pseudoannulata as a potential eco-friendly predator against pests.  相似文献   

18.
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.  相似文献   

19.
1. It is increasingly realised that aquatic and terrestrial systems are closely linked. We investigated stable isotope variations in Odonata species, putative prey and basal resources of aquatic and terrestrial systems of northern Mongolia during summer. 2. In permanent ponds, δ13C values of Odonata larvae were distinctly lower than those of putative prey, suggesting that body tissue comprised largely of carbon originating from isotopically light carbon sources. Presumably, prey consumed during autumn and winter when carbon is internally recycled and/or methanotrophic bacteria form an important basal resource of the food web. In contrast, in a temporary pond, δ13C values of Odonata larvae were similar to those of putative prey, indicating that their body carbon originated mainly from prey species present. 3. Changes in δ15N and δ13C values between larvae and adults were species specific and reflected differential replacement of the larval isotopic signature by the terrestrial diet of adult Odonata. The replacement was more pronounced in Odonata species of permanent ponds than in those of the temporary pond, where larvae hatched later in the year. Replacement of larval carbon varied between tissues, with wings representing the larval isotopic signature whereas thoracic muscles and eggs reflected the δ15N and δ13C values of the terrestrial diet of adults. 4. The results suggest that because of their long larval development, Odonata species of permanent ponds carry the larval signature, which is partly replaced during their terrestrial life. Terrestrial prey forms the basis for egg production and thus the next generation of aquatic larvae. In temporary ponds, in contrast, Odonata species rely on prey from a single season, engage in a prolonged aquatic phase and hatch later, leaving less time to acquire terrestrial prey resources for offspring production. Stable isotope analysis provided important insights into the food webs of the waterbodies and their relationship to the terrestrial system.  相似文献   

20.
  1. Generalistic interactions between predator and prey may vary with ecosystem type, predator traits, and prey traits, but the interplay of these factors has not been assessed in ground food webs.
  2. We investigated trophic interactions of ground-dwelling spiders across eight forests in European Russia associated with body size, hunting strategy, microhabitat specialization, potential prey type, potential prey population density, and forest type (coniferous vs. broadleaved). We analyzed 128 individual spiders, including juveniles, all identified to the family level with two complementary methods: molecular gut content analysis, and stable isotope analysis of carbon and nitrogen.
  3. The results suggest that feeding frequency of spiders is affected by predator body size and by selection of certain prey type. Stable isotope analysis showed similar trophic niches among spider families, varying moderately with forest type. Larger spiders had higher Δ13C values than smaller ones, but similar Δ15N values, suggesting that different size classes of spiders belong to different food chains. Results based on stable isotope and molecular gut content analyses were weakly linked, indicating them targeting different trophic niche dimensions.
  4. At least for the group-level interactions, family identity and hunting strategy of predator has little predictive power while predator body size and prey traits affected trophic niche dimensions calling for future studies in this direction. Large spiders feed more and rely on different basal resources than small spiders, suggesting that including small species and juveniles provides a more comprehensive picture of food web organization.
  相似文献   

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