Phytophagous insects enhance nitrogen flux in a desert creosotebush community

Summary We tested the hypothesis that herbivorous insects on desert shrubs contribute to short-term nitrogen cycling, and increase rates of nitrogen flux from nutrient rich plants. Creosotebush (Larrea tridentata) shrubs were treated with different combinations of fertilizer and water augmentations, resulting in different levels of foliage production and foliar nitrogen contents. Foliage arthropod populations, and nitrogen in canopy dry throughfall, wet throughfall and stemflow were measured to assess nitrogen flux rates relative to arthropod abundances on manipulated and unmanipulated shrubs over a one-month period during peak productivity. Numbers and biomass of foliage arthropods were significantly higher on fertilized shrubs. Sap-sucking phytophagous insects accounted for the greatest numbers of foliage arthropods, but leaf-chewing phytophagous insects represented the greatest biomass of foliage arthropods. Measured amounts of bulk frass (from leaf-chewing insects) were not significantly different among the various treatments. Amounts of nitrogen from dry and wet throughfall and stemflow were significantly greater under fertilized shrubs due to fine frass input from sap-sucking insects. Increased numbers and biomass of phytophagous insects on fertilized shrubs increased canopy to soil nitrogen flux due to increased levels of herbivory and excrement. Nitrogen excreted by foliage arthropods accounted for about 20% of the total one month canopy to soil nitrogen flux, while leaf litter accounted for about 80%.     

Tracking nutrients in space and time: Interactions between grazing lawns and drought drive abundances of tallgrass prairie grasshoppers

1. We contrast the response of arthropod abundance and composition to bison grazing lawns during a drought and non‐drought year, with an emphasis on acridid grasshoppers, an important grassland herbivore.
2. Grazing lawns are grassland areas where regular grazing by mammalian herbivores creates patches of short‐statured, high nutrient vegetation. Grazing lawns are predictable microsites that modify microclimate, plant structure, community composition, and nutrient availability, with likely repercussions for arthropod communities.
3. One year of our study occurred during an extreme drought. Drought mimics some of the effects of mammalian grazers: decreasing above‐ground plant biomass while increasing plant foliar percentage nitrogen.
4. We sampled arthropods and nutrient availability on and nearby (“off”) 10 bison‐grazed grazing lawns in a tallgrass prairie in NE Kansas. Total grasshopper abundance was higher on grazing lawns and the magnitude of this difference increased in the wetter year of 2019 compared to 2018, when drought led to high grass foliar nitrogen concentrations on and off grazing lawns. Mixed‐feeding grasshopper abundances were consistently higher on grazing lawns while grass‐feeder and forb‐feeder abundances were higher on lawns only in 2019, the wetter year. In contrast, the abundance of other arthropods (e.g., Hemiptera, Hymenoptera, and Araneae) did not differ on and off lawns, but increased overall in 2019, relative to the drought of 2018.
5. Understanding these local scale patterns of abundances and community composition improves predictability of arthropod responses to ongoing habitat change.

     

Arctic arthropod assemblages in habitats of differing shrub dominance

Recent climate warming in the Arctic has caused advancement in the timing of snowmelt and expansion of shrubs into open tundra. Such an altered climate may directly and indirectly (via effects on vegetation) affect arctic arthropod abundance, diversity and assemblage taxonomic composition. To allow better predictions about how climate changes may affect these organisms, we compared arthropod assemblages between open and shrub‐dominated tundra at three field sites in northern Alaska that encompass a range of shrub communities. Over ten weeks of sampling in 2011, pitfall traps captured significantly more arthropods in shrub plots than open tundra plots at two of the three sites. Furthermore, taxonomic richness and diversity were significantly greater in shrub plots than open tundra plots, although this pattern was site‐specific as well. Patterns of abundance within the five most abundant arthropod orders differed, with spiders (Order: Araneae) more abundant in open tundra habitats and true bugs (Order: Hemiptera), flies (Order: Diptera), and wasps and bees (Order: Hymenoptera) more abundant in shrub‐dominated habitats. Few strong relationships were found between vegetation and environmental variables and arthropod abundance; however, lichen cover seemed to be important for the overall abundance of arthropods. Some arthropod orders showed significant relationships with other vegetation variables, including maximum shrub height (Coleoptera) and foliar canopy cover (Diptera). As climate warming continues over the coming decades, and with further shrub expansion likely to occur, changes in arthropod abundance, richness, and diversity associated with shrub‐dominated habitat may have important ecological effects on arctic food webs since arthropods play important ecological roles in the tundra, including in decomposition and trophic interactions.     

Seasonal Distribution and Diversity of Ground Arthropods in Microhabitats Following a Shrub Plantation Age Sequence in Desertified Steppe

In desertified regions, shrub-dominated patches are important microhabitats for ground arthropod assemblages. As shrub age increases, soil, vegetation and microbiological properties can change remarkably and spontaneously across seasons. However, relatively few studies have analyzed how ground arthropods respond to the microhabitats created by shrubs of different plantation ages across seasons. Using 6, 15, 24 and 36 year-old plantations of re-vegetated shrubs (Caragana koushinskii) in the desert steppe of northwestern China as a model system, we sampled ground arthropod communities using a pitfall trapping method in the microhabitats under shrubs and in the open areas between shrubs, during the spring, summer and autumn. The total ground arthropod assemblage was dominated by Carabidae, Melolonthidae, Curculionidae, Tenebrionidae and Formicidae that were affected by plantation age, seasonal changes, or the interaction between these factors, with the later two groups also influenced by microhabitat. Overall, a facilitative effect was observed, with more arthropods and a greater diversity found under shrubs as compared to open areas, but this was markedly affected by seasonal changes. There was a high degree of similarity in arthropod assemblages and diversity between microhabitats in summer and autumn. Shrub plantation age significantly influenced the distribution of the most abundant groups, and also the diversity indices of the ground arthropods. However, there was not an overall positive relationship between shrub age and arthropod abundance, richness or diversity index. The influence of plantation age on arthropod communities was also affected by seasonal changes. From spring through summer to autumn, community indices of ground arthropods tended to decline, and a high degree of similarity in these indices (with fluctuation) was observed among different ages of shrub plantation in autumn. Altogether the recovery of arthropod communities was markedly affected by seasonal variability, and they demonstrated distinctive communal fingerprints in different microhabitats for each plantation age stage.     

Biotic filtering and mass effects in small shrub patches: is arthropod community structure predictable based on the quality of the vegetation?

1. Community assembly is affected by four processes: dispersal, filtering effects (selection), ecological drift and evolution. The role of filtering relative to dispersal and drift should decline with patch size, hampering possibilities to predict which organisms will be observed within small‐sized patches. However, vegetation structure is known to have a marked impact on species assemblages, and plant quality may act as a biotic filter. This challenges the assumption of unpredictable species assemblages in small‐sized vegetation patches. 2. Using 32 stands of five shrub species in south‐west Finland, this study investigated whether biotic filtering effects caused by patch‐forming plants are strong enough to overcome the mixing of mobile arthropod assemblages across small patches. 3. Stochastic variation did not hide the signals of biotic filtering and dispersal in the small shrub patches. Habitat richness around the patches explained a three times larger share of variation in the species composition than did the identity of the patch‐forming plant, but it had less effect on the abundance of arthropods. A radius of 50–100 m around a patch explained the species composition best. 4. Abundance patterns varied between the feeding guilds; the patch‐forming shrub influenced the abundances of detritivores and leaf‐feeding herbivores, whereas the abundances of flower‐visiting herbivores appeared to track the flowering phenology of the plants. Shrub identity had little effect on omnivores or predators. Predator abundances were correlated with the abundance of potential prey. 5. The results of this study suggest that community composition within a vegetation patch may be predictable even if dispersal overrides local filtering effects, as suggested by the mass‐effects paradigm.     

Shrub shading moderates the effects of weather on arthropod activity in arctic tundra

1. Rapid warming has facilitated an increase in deciduous shrub cover in arctic tundra. Because shrubs create a cooler microclimate during the growing season, shrub cover could modulate the effects of global warming on the phenology and activity of ectotherms, including arthropods. This possibility was explored here using two dominant arthropod groups (flies and wolf spiders) in Alaskan tundra. 2. We monitored arthropods with pitfall traps over five summers at four sites that differed in shrub abundance, and used generalised additive mixed models (GAMMs) to separate the two underlying components of pitfall trap catch: the seasonal trend in arthropod density and the effects of short‐term weather variation (air temperature, wind speed, rainfall, solar radiation) on arthropod activity. 3. We found that shrub cover significantly altered the seasonal trend in the abundance of flies by reducing early‐season pitfall catch, in line with observed later snowmelt in shrub‐dominated plots at these sites. 4. Additionally, shrub cover modulated the effects of many weather variables on arthropod activity: shrub cover shifted wolf spiders' temperature–activity relationship, dampened the positive effect of solar radiation on the activity of arthropods in total, and ameliorated the negative effect of wind on the activity of flies. 5. Thus, these results indicate that shrub encroachment will probably be accompanied by altered arthropod responses to warming and other key weather variables. Because the rate of key ecological processes – herbivory, decomposition, predation – are controlled by activity at the organismal level, these effects on arthropods will have long‐term ecosystem‐level consequences.     

Water stress,leaf nutrients and defoliation: a model of dieback of rural eucalypts

Water stress, concentrations of foliar nutrients and damage of foliage by insects were studied over an eleven month period, for eight dieback eucalypts and eight closely matched healthy trees growing in close association on two grazing properties near Brisbane. Four healthy eucalypts in a neighbouring State Forest were also studied. The study region had suffered severe climatic stress between 1972 and 1976 when dieback was first observed by residents and when high populations of defoliating insects were observed. Dieback trees had a higher proportion of foliage damaged by insects and higher concentrations of foliar nutrients than did matched healthy trees. The study period was one of low climatic stress during which dieback and healthy trees followed similar seasonal patterns of predawn xylem pressure potential. Dieback trees developed lower daytime minimum xylem pressure potentials than did matched healthy trees, and differed in their stomatal responses. A model of initiation and development of rural eucalypt dieback is proposed. This may have general application to many non-specific rural diebacks in which heavy insect damage is implicated. The balance between rural eucalypts and their insect herbivores is precarious. Any factor capable of causing extensive defoliation, or an increase in foliar nitrogen, or an increase in populations of insect herbivores may upset this balance. A positive feedback loop may be activated, whereby the production of nitrogen rich epicormic foliage enhances a build up of insect populations. Repeated insect defoliation leads to tree dieback. Evidence in the literature supporting the model is reviewed and aspects requiring further research are outlined.     

Variation in foliar nutrients in Eucalyptus trees in eastern and Western Australia

Abstract Levels of nitrogen, phosphorus and potassium were measured for the foliage of two co-dominant eucalypts at each of two sites, one in eastern Australia and the other in Western Australia. In eastern Australia, foliage was sampled in the canopy and subcanopy for narrow-leaved ironbark Eucalyptus crebra and grey box E. mollucana and in Western Australia, for jarrah E. marginata and marri E. calophylla. The Western Australian trees were also sampled for ‘young’ and ‘old’ leaves. Both eucalypts in eastern Australia had greater nitrogen and phosphorus levels, but lower potassium, than E. marginata or E. calophylla. Eucalyptus calophylla foliage had greater levels of all three nutrients than E. marginata foliage as did E. crebra relative to E. mollucana. At both sites, foliar nutrient levels were greater in the canopy than subcanopy foliage, and, at least in Western Australia, the younger leaves had greater nutrient levels than the older leaves. The observed differences in foliar nutrient levels are consistent with observed trends in the abundance and diversity of foliage arthropods and the use of the trees as foraging substrates by birds.     

More is not necessarily better: the impact of limiting and excessive nutrients on herbivore population growth rates

Abstract 1. The body tissues of insect herbivores contain higher concentrations of nitrogen and phosphorus than do their host plants, leading to an elemental mismatch that can limit herbivore growth, fecundity and ultimately influence population dynamics. While low nutrient availability can limit herbivore growth and reproduction, nutrient levels that exceed an organism’s nutritional requirements, i.e. an organisms’ threshold elemental ratio, can also decrease performance. 2. We conducted a laboratory experiment to examine the impacts of nitrogen and phosphorus additions on population growth rates of a phloem‐feeding insect herbivore. 3. Herbivore per capita population growth rates were highest at intermediate foliar nitrogen concentrations, indicating a performance cost on the highest nitrogen foliage. While there was no direct effect of foliar phosphorus concentration on insect performance, there was a strong and unexpected indirect effect. High soil phosphorus availability increased both foliar nitrogen concentrations and aphid tissue nitrogen, resulting in low population growth rates when both soil nitrogen and phosphorus availabilities were high. 4. In this study, experimental increases in foliar nitrogen levels led to a decrease in herbivore performance suggesting that excessive nutrient levels can limit herbivore population growth rates.     

Nutrient stress: an explanation for plant anti-herbivore responses to defoliation

Summary A hypothesis is put forward that the long-lasting inducible responses of trees to herbivores, particularly lepidopteran defoliators, may not be active defensive responses, but a by-product of mechanisms which rearrange the plant carbon/nutrient balance in response to nutrient stress caused by defoliation. When defoliation removes the foliage nutrients of trees growing in nutrient-poor soils, it increases nutrient stress wich in turn results in a high production of carbon-based allelochemicals. The excess of carbon that cannot be diverted to growth due to nutrient stress is diverted to the production of plant secondary metabolites. The level of carbon-based secondary substances decays gradually depending on the rate at which nutrient stress is relaxed after defoliation. In nutrient-poor soils and in plant species with slow compensatory nutrient uptake rates the responses induced by defoliation can have relaxation times of several years. The changes in leaf nitrogen and phenolic content of mountain birch support this nutrient stress hypothesis. Defoliation reduces leaf nitrogen content while phenolic content increases. These responses of mountain birch to defoliation are relaxed within 3–4 years.     

Intercropping in high input agriculture supports arthropod diversity without risking significant yield losses

Arthropod diversity of different taxonomic groups and ecosystem services are declining, yet current measures to counteract losses are often restricted to small areas of land or field margins, particularly in agricultural systems. At the same time, large areas of land will be required to feed a growing global population. Intercropping has been proposed as a potential solution to maximize both biodiversity and yield at large scale, but experimental evidence is scarce.In a three-year field experiment, we manipulated crop diversity and management intensity in a cereal-legume intercropping experiment in Germany, where 50% of wheat was replaced by faba beans. We measured arthropod abundance and diversity of different functional groups (pollinators, natural enemies, herbivores) and crop yield.We found that increasing crop diversity increased abundance and diversity of arthropods. Notably, pollinator and natural enemy abundances increased in intercropped systems. Low management intensity generally had positive effects on arthropod abundance and especially on pollinator diversity, indicating benefits of reduced inputs of fertilizers and herbicides. While wheat yield was higher in monocultures and for high management intensity, total grain yield of the intercrop (indicated by land equivalent ratio) was higher in mixtures. We found that trade-offs were stronger between arthropod diversity and wheat yield than between arthropod abundance and wheat yield. Specialist wheat herbivores and generalist herbivores were more abundant at higher wheat yields. Conversely, pollinator and natural enemy diversity were negatively associated with wheat yield.Our results show that diversification can promote both higher yields and greater diversity of arthropods. Intercropping can thus be an opportunity to support biodiversity without risking significant yield losses.     

干旱区不同年龄灌丛斑块地面节肢动物的聚集效

以荒漠草原区6、15、24、36年生柠条灌丛为研究对象,调查了不同林龄灌丛内外土壤性质和地面节肢动物群落特征,分析了干旱区灌丛斑块生境中地面节肢动物群落随灌木林发育的变化特征.结果表明： 1)6年生林地只对土壤物理性质(土壤质地、含水量、pH和电导率)产生显著影响,从15年开始林地土壤有机质和养分(N、P)显著增加.2)在研究样地共捕获27个节肢动物类群,步甲科、拟步甲科和蚁科是优势类群.从6到15年生林地,地面节肢动物优势类群数减少,常见类群数增加.从15年生经24年生到36年生林地,灌丛内外类群组成差异先缩小后变大.36年生林地某些特殊的动物类群开始大量出现,如蜣螂在36年生林地灌丛中成为优势类群.3)6和24年生林地灌丛内外地面节肢动物群落结构差别较小,15和36年生林地灌丛内外地面节肢动物群落结构差异显著.4)林龄对灌丛内地面节肢动物的分布影响较大,而对灌丛外地面节肢动物的分布影响较小.灌丛发育过程通过对灌丛下微生境的改变如土壤质地、电导率、pH等来显著影响地面节肢动物的分布,并对灌丛外样地产生一定的辐射作用.干旱区灌丛发育过程对地面节肢动物聚集效应产生显著影响,在不同林龄阶段这种聚集效应表现不一,直接影响沙化草地生态系统的恢复过程.     

Greater shrub dominance alters breeding habitat and food resources for migratory songbirds in Alaskan arctic tundra

Climate warming is affecting the Arctic in multiple ways, including via increased dominance of deciduous shrubs. Although many studies have focused on how this vegetation shift is altering nutrient cycling and energy balance, few have explicitly considered effects on tundra fauna, such as the millions of migratory songbirds that breed in northern regions every year. To understand how increasing deciduous shrub dominance may alter breeding songbird habitat, we quantified vegetation and arthropod community characteristics in both graminoid and shrub dominated tundra. We combined measurements of preferred nest site characteristics for Lapland longspurs (Calcarius lapponicus) and Gambel's White‐crowned sparrows (Zonotrichia leucophrys gambelii) with modeled predictions for the distribution of plant community types in the Alaskan arctic foothills region for the year 2050. Lapland longspur nests were found in sedge‐dominated tussock tundra where shrub height does not exceed 20 cm, whereas White‐crowned sparrows nested only under shrubs between 20 cm and 1 m in height, with no preference for shrub species. Shrub canopies had higher canopy‐dwelling arthropod availability (i.e. small flies and spiders) but lower ground‐dwelling arthropod availability (i.e. large spiders and beetles). Since flies are the birds' preferred prey, increasing shrubs may result in a net enhancement in preferred prey availability. Acknowledging the coarse resolution of existing tundra vegetation models, we predict that by 2050 there will be a northward shift in current White‐crowned sparrow habitat range and a 20–60% increase in their preferred habitat extent, while Lapland longspur habitat extent will be equivalently reduced. Our findings can be used to make first approximations of future habitat change for species with similar nesting requirements. However, we contend that as exemplified by this study's findings, existing tundra modeling tools cannot yet simulate the fine‐scale habitat characteristics that are critical to accurately predicting future habitat extent for many wildlife species.     

Changes in arthropod diversity along a land use driven gradient of shrub cover in savanna rangelands: identification of suitable indicators

Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground-dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.     

The seasonal dynamics of leaf resin,nitrogen, and herbivore damage in Eriodictyon californicum and their parallels in Diplacus aurantiacus

Summary The chaparral shrub Eriodictyon californicum produces a flavonoid leaf resin with a chemically similar composition to that previously reported for the sympatric shrub Diplacus aurantiacus. We determined the phenology, resin content, and nitrogen content of Eriodictyon leaves and the leaf area lost to herbivores. Nitrogen content and resin content were both negatively correlated with leaf age at each sampling date, but nitrogen decreased during the growing season while resin increased. The fraction of leaf area lost during the growing season averaged less than 7% and was highest on the oldest leaves. The seasonal pattern of resin production in Eriodictyon corresponds to that in Diplacus, indicating that the similar environments of Eriodyctyon and Diplacus have led to convergent leaf resins. This convergence in these two plants has implications for chemicals of similar function in other chaparral shrubs.     

Effect of fragmentation on predation pressure of insect herbivores in a north temperate deciduous forest ecosystem

1. Forest fragmentation affects many ecosystem processes by spatially altering relationships among organisms. Herbivory by arthropods is an important ecosystem processes in forests that fragmentation alters by changing relationships among herbivores, their predators, and their hosts. The relative importance of these factors remains unclear. 2. It was tested whether the exclusion of vertebrate predators affected the arthropod abundance or amounts of herbivory in a fragmented, deciduous forest landscape in southern Quebec. Differences in the abundance of arthropod herbivores and amounts of herbivory in forest patches with different landscape characteristics (small isolated patches versus large connected ones), on sugar maple saplings with or without exclosures that restricted access by large vertebrate predators were measured. 3. Saplings protected from predators with exclosures had greater abundances of all arthropods (herbivores and invertebrate predators) than those without, indicating potential top‐down effects of vertebrate predators on arthropods. Analysis of effect sizes between exclosure treatments and controls suggests the magnitude of predation effects may be affected by fragmentation. 4. Strong top‐down effects of predators on arthropods, and weak effects of fragmentation on predation or amounts of herbivory were found. As a result, herbivory may be regulated by factors other than vertebrate predation in this system.     

Influence of leaf traits on the spatial distribution of arboreal arthropods within an overstorey rainforest tree

Abstract.

• 1 Several attributes of foliage were measured from the Australian rainforest tree Argyrodendron actinophyllum Edlin (Sterculiaceae). These were related to estimates of abundance per leaf area of the most common arthropod guilds and families sampled with restricted canopy fogging.
• 2 When all these arthropod groups were considered, much of the overall variance in arthropod spatial distribution could be attributed to leaf age characteristics, arthropod aggregation patterns, arthropod activity and distance to tree trunk.
• 3 The fraction of variance which could be specifically explained by foliage attributes such as nitrogen-, water- and fibre-content, specific leaf weight, and epiphyll load was small for most arthropod groups (usually <30%). However, an index of food quality explained a higher proportion of variance (50%) in the abundance of phloem-feeders. Leaf size and foliage compactness did not influence significantly the abundance of any arthropod group.
• 4 Most herbivores were more abundant on young foliage than on mature leaves. With the exception of Corylophidae and Chrysomelidae, which were more abundant in the lower and upper canopy respectively, arthropod stratification was not conspicuous within the inner core of tree crowns.
• 5 The results firstly emphasize the distribution of young foliage as a key factor affecting the abundance of many herbivores and, secondly, the importance of the local illumination regime for host leaf production and its indirect effects on the spatial distribution of arboreal arthropods.

     

Soil nutrients under shrub hummocks and debris mounds two decades after ploughing

Aims

Shrub removal by ploughing has been used widely to reduce the effects of shrub encroachment into open woodlands and grasslands. Our aim was to demonstrate that soil chemical properties varied markedly among three patch types (shrub hummock, debris mound, interspace) which varied in age, almost two decades after shrub removal by ploughing.

Methods

We compared changes in nutrients under 1) young post-ploughing recruits and mature, unploughed shrubs, 2) mature and recently formed debris mounds and 3) ploughed and recovering interspaces at three depths.

Results

Irrespective of their age, nutrient concentrations were greater under shrub hummocks and debris mounds than in the interspaces at two sites. Soil in mature shrub hummocks generally had greater levels of labile carbon and nitrogen (total, mineral, mineralisable), but results varied between sites. There were a few, sometimes inconsistent, effects of ploughing on nutrients under debris mounds, and no differences between the interspaces two decades after ploughing. Nutrient effects were most marked in the top 15 cm of the soil, diminishing rapidly with depth.

Conclusions

Our results reinforce the importance of hummocks and mounds as resource sinks and indicate the long-lasting effects of disturbances such as ploughing on soil nutrient pools.     

Arthropod food webs become increasingly lipid‐limited at higher trophic levels

Understanding why food chains are relatively short in length has been an area of research and debate for decades. We tested if progressive changes in the nutritional content of arthropods with trophic position limit the availability of a key nutrient, lipid, for carnivores. Arthropods at higher trophic levels had progressively less lipid and more protein in their bodies compared with arthropods at lower trophic levels. The nutrients present in arthropod bodies were directly related to the nutrients that predators extracted when feeding on those arthropods. As a consequence, nutrient assimilation shifted from lipid‐biased to protein‐biased as arthropods ascended trophic levels from herbivores to secondary carnivores. Successive changes in the nutritional consequences of predation may ultimately set an upper limit on the number of trophic levels in arthropod communities. Further work is needed to examine the influence of lipid and other nutrients on food web traits in a range of ecosystems.     

Abundance and stratification of foliage arthropods in a lowland rain forest of Cameroon

Abstract.

• 1 Arthropod densities and apparent leaf damage were compared within the canopy ecotone and the shrub layer of a lowland rain forest in Cameroon, using a branch clipping method.
• 2 Most of the individuals collected consisted of ants (average 44%) and various herbivores (31%). Overall arthropod densities amounted to 17 individuals per sample, which, on average, consisted of 0.85 m² of foliage area. Arthropod densities were lower than on temperate foliage.
• 3 Arthropod densities were about 3 times higher in the canopy than within the shrub layer. In particular, ants and herbivores were significantly more abundant in the canopy than within the shrub layer. Usually, layer effects rather than site effects appeared to cause greater variance in arthropod abundance.
• 4 Arthropod species-richness, as estimated by the number of operational taxonomic units sorted, was higher in canopy samples than in samples obtained from the shrub layer. However, apparent leaf damage was higher within the shrub layer (10.9%) than on the canopy (5.2%).
• 5 Possible factors responsible for the high densities of ants and herbivores on the canopy and for the high leaf damage within the shrub layer are discussed.