Individual tree context and contrast dictate tree physiological features and arthropod biodiversity patterns across multiple trophic levels

1. The present study tested the hypothesis that tree context (natural, semi-natural or planted) and contrast (height of surrounding vegetation) affect tree physiological characters (leaf size, nutrient content, and stress-related factors), and also alter the arthropod biodiversity patterns either directly or indirectly. 2. Arthropods were collected from tree canopies using chemical fogging from the ecologically important South African native tree Podocarpus elongatus. 3. Low contrast trees had significantly larger leaves than those in high contrast environments and harboured an overall higher richness, abundance, and biomass of arthropods, although this was guild-dependent. 4. Trees in natural contexts had less foliar δ¹³C, suffered less from artificial nitrogen enrichment, and harboured significantly higher herbivore and predator arthropod richness and unique assemblages, compared with planted trees. 5. Semi-natural trees supported natural levels of arthropod richness, although these were mostly generalist species. 6. Tree context and contrast can therefore dictate associated biota at multiple trophic levels and native trees may fail to maintain natural biodiversity in transformed landscapes.     

Consumer diversity mediates invasion dynamics at multiple trophic levels

Theory and recent experiments, mostly focused on plants, indicate that biodiversity can reduce invasion success, but diversity effects on mobile animal invasion have received little attention. We tested effects of mobile crustacean grazer diversity (species richness) on the establishment of invaders at multiple trophic levels in flow-through seagrass mesocosms. On average, increasing diversity of resident grazers reduced population growth and biomass of experimentally introduced grazers. This increase in invasion resistance was concurrent with reductions in food and habitat availability and increases in resident density, paralleling previous results with plants. In many cases, mixtures of resident species resisted invasion better than did any single resident species, arguing that interactions among residents, rather than a sampling mechanism, explained diversity effects on invasion. Higher grazer diversity also generally reduced biomass of naturally recruiting invertebrates and algae and shifted epiphytic community dominance from algae to sessile invertebrates. Exploitation competition, then, appears to contribute to the diversity effect on invasion in both plant and animal systems. Our results further suggest that resident competitive advantage may also be at work in multi-trophic level systems. Thus, negative effects of local diversity on invasion appear general, and trophically mediated processes can also strongly influence invader success and identity in multi-trophic level systems.     

Species richness changes across two trophic levels simultaneously affect prey and consumer biomass

Increasing species richness of primary producers or consumers is proposed to increase primary and secondary production; however, the consequences of biodiversity change across trophic levels has been poorly investigated. We used a controlled marine microbial system to investigate the effects of simultaneous changes in biodiversity of consumer and prey species. Consumer (ciliates) and prey (algae) richness and identity were manipulated independently in a complete factorial design. The results showed clear biodiversity effects of both consumers and prey, within and across trophic levels. We found reduced prey and increased consumer biomass with increased consumer richness, with the most diverse prey assemblage supporting the highest biomass of consumers at the highest richness of consumers. Increasing prey richness did not increase resistance to consumption when consumers were present. Instead, our results indicated enhanced energy transfer with simultaneous increasing richness of consumers and prey.     

Effects of plant sex on insect abundance across three trophic levels in the perennial shrub Buddleja cordata

Although there is a growing interest in the effects of intra‐specific plant genetic variation on species interactions, the effects of plant sex, an important axis of genetic variation, have been less studied. In addition, previous work investigating plant sex effects on species interactions has frequently focused on bitrophic interactions (e.g., herbivory), usually ignoring plant sex effects on higher trophic levels (i.e., natural enemies). Here, we investigated the effects of plant sex on herbivore abundance and that of their natural enemies associated with the dioecious shrub Buddleja cordata Kunth (Scrophulariaceae). Furthermore, we measured a subset of plant traits frequently involved in herbivore resistance and the potentially underlying plant sex effects. To this end, we recorded the abundances of a specialist leaf‐chewing caterpillar [Acronyctodes mexicanaria Walker (Lepidoptera: Geometridae)] throughout an entire growing season. We also recorded information about the caterpillar’s parasitoids, as well as leaf water content, phenolic compounds, phosphorus, and nitrogen for male and female plants of B. cordata. Plant sex did not significantly influence caterpillar abundance but did have an effect on natural enemies, with parasitoid abundance being 2.4‐fold greater on female than on male plants. The effect of plant sex on parasitoids remained significant after accounting for caterpillar abundance, suggesting that it was underlain by a trait‐mediated (rather than density‐mediated) mechanism. Finally, we found that male plants had a higher concentration of phenolic compounds (other traits did not differ between plant sexes). These results provide valuable evidence for the extended effects of plant sex on the third trophic level and point at plant traits potentially mediating such effects.     

Plant and insect diversity along a pollution gradient: understanding species richness across trophic levels

We analysed species richness of plants and true bugs (Insecta, Heteroptera) along a pollution gradient in Scots pine stands in Central Germany. As a consequence of particulate deposition, pH-values of soils increased in the vicinity of the emission source. Therefore, emission increased productivity. Species richness of plants increased with decreasing distance from emission source, and thus with increasing productivity. Similarly, species richness of herbivorous Heteroptera increased with decreasing distance from emission source, whereas, surprisingly, abundance decreased. The proportion of specialised herbivorous bug species is largest in the vicinity of the emission source. Thus, the diversity pattern of herbivores may be explained by the specialisation hypothesis and not the consumer rarity hypothesis. Species richness and abundance of carnivorous Heteroptera showed no significant trend along the gradient. Overall our data favour the bottom-up control of species diversity in the analysed system.     

Pyramids of species richness: the determinants and distribution of species diversity across trophic levels

How species richness is distributed across trophic levels determines several dimensions of ecosystem functioning, including herbivory, predation, and decomposition rates. We perform a meta‐analysis of 72 large published food webs to investigate their trophic diversity structure and possible endogenous, exogenous, and methodological causal variables. Consistent with classic theory, we found that published food webs can generally be described as ‘pyramids of species richness’. The food webs were more predator‐poor, prey‐rich and hierarchical than is expected by chance or by the niche or cascade models. The trophic species richness distribution also depended on centrality, latitude, ecosystem‐type and methodological bias. Although trophic diversity structure is generally pyramidal, under many conditions the structure is consistently uniform or inverse‐pyramidal. Our meta‐analysis adds nuance to classic assumptions about food web structure: diversity decreases with trophic level, but not under all conditions, and the decrease may be scale‐dependent. Synthesis The distribution of species richness across trophic levels has not been evaluated in recent decades, despite improvement in food web resolution and the relevance of biodiversity distribution to ecosystem function. Our meta‐analysis of 72 large, recent food webs, illustrates that published food webs can generally be described as basal‐rich, top‐poor ‘pyramids of species richness’, consistent with classic theory. Although trophic diversity structure is generally pyramidal, under some environmental and ecological conditions the structure is uniform or inverse‐pyramidal. Our meta‐analysis confirms classic theory about food web structure, while adding nuance by describing conditions under which classic pyramid structure is not observed.     

Predicting effects of multiple interacting global change drivers across trophic levels

Global change encompasses many co-occurring anthropogenic drivers, which can act synergistically or antagonistically on ecological systems. Predicting how different global change drivers simultaneously contribute to observed biodiversity change is a key challenge for ecology and conservation. However, we lack the mechanistic understanding of how multiple global change drivers influence the vital rates of multiple interacting species. We propose that reaction norms, the relationships between a driver and vital rates like growth, mortality, and consumption, provide insights to the underlying mechanisms of community responses to multiple drivers. Understanding how multiple drivers interact to affect demographic rates using a reaction-norm perspective can improve our ability to make predictions of interactions at higher levels of organization—that is, community and food web. Building on the framework of consumer–resource interactions and widely studied thermal performance curves, we illustrate how joint driver impacts can be scaled up from the population to the community level. A simple proof-of-concept model demonstrates how reaction norms of vital rates predict the prevalence of driver interactions at the community level. A literature search suggests that our proposed approach is not yet used in multiple driver research. We outline how realistic response surfaces (i.e., multidimensional reaction norms) can be inferred by parametric and nonparametric approaches. Response surfaces have the potential to strengthen our understanding of how multiple drivers affect communities as well as improve our ability to predict when interactive effects emerge, two of the major challenges of ecology today.     

Earthworms promote greater richness and abundance in the emergence of plant species across a grassland-forest ecotone

Aims Chalk grasslands are subject to vegetation dynamics that range from species-rich open grasslands to tall and encroached grasslands, and woods and forests. In grasslands, earthworms impact plant communities and ecosystem functioning through the modification of soil physical, chemical and microbiological properties, but also through their selective ingestion and vertical transportation of seeds from the soil seed bank. Laboratory experiments showed that seed–earthworm interactions are species specific, but little is known on the impact of seed–earthworm interactions in the field. The overall aim of this study was to better understand seed–earthworm interactions and their impact on the plant community. First we analyzed the composition of seedlings emerging from casts after earthworm ingestion. Then we compared seedling composition in casts to the plant composition of emerging seedlings from the soil and of the aboveground vegetation along four stages of the secondary succession of chalk grasslands.Methods Four stages of the secondary succession of a chalk grassland—from open sward to woods—were sampled in Upper Normandy, France, in February 2010. Within each successional stage (×3 replicates), we sampled the standing vegetation, soil seed bank at three soil depths (0–2, 2–5 and 5–10cm) and earthworm surface casts along transects. Soil and cast samples were water sieved before samples were spread onto trays and placed into a greenhouse. Emerging seedlings were counted and identified. Effect of successional stage and origin of samples on mean and variability of abundance and species richness of seedlings emerging from casts and soil seed banks were analyzed. Plant compositions were compared between all sample types. We used generalized mixed-effect models and a distance-based redundancy multivariate analysis.Important findings Seedling abundance was always higher in earthworm casts than in the soil seed bank and increased up to 5-fold, 4-fold and 3.5-fold, respectively, in the tall grassland, woods and encroached grassland compared to the soil surface layer. Species richness was also higher in earthworm casts than in the soil seed bank in all successional stages, with a 4-fold increase in the encroached grassland. The plant composition of the standing vegetation was more similar to that of seedlings from casts than to that of seedlings from the soil seed bank. Seedlings diversity emerging from casts in the tall and encroached grasslands tended toward the diversity found in woods. Our results indicate that earthworms may promote the emergence of seedlings. We also suggest that the loss of some plant species in the seed bank and the tall grass vegetation in intermediary successional stages modify the local conditions and prevent the further establishment of early-successional plant species.     

互利关系和植物多样性对节肢动物群落不同营养级的影响路径和强度

食物网中的上行效应和下行效应对于群落的动态和生态系统功能有十分重要的影响,旨在探讨互利关系和植物多样性对节肢动物群落中食物网不同营养级之间的影响。通过随机裂区试验方法,分别设置了3种蚂蚁-紫胶虫互利关系处理(有互利关系、无互利关系和自然对照)以及3种植物多样性处理(单一种植、2树种混植和3树种混植),于2016年8月和9月分两次用手捡法、网扫法和震落法采集试验地寄主植物上所有的节肢动物,并按照不同营养级将其分类。利用结构方程模型分析方法对不同营养级之间的相互作用的路径和强度进行了比较,结果显示:1)互利关系对捕食者和消费者均有显著的下行作用,有互利关系处理下蚂蚁对捕食者的路径强度要强于自然对照组,互利关系对捕食者的影响要强于对消费者的影响。2)植物多样性会通过影响植物的生物量而对消费者和捕食者产生显著的上行效应影响,这种影响会随着营养级的升高而显著减小。3)消费者主要受植物多样性的上行效应影响,而捕食者主要受互利关系的下行效应影响。有互利关系的食物网结构更加复杂,营养级之间的相互作用更为显著。探讨了以蚂蚁-紫胶虫互利关系为核心作用的紫胶林生态系统中互利关系和植物多样性对节肢动物食物网中...     

Fish diversity as a determinant of ecosystem properties across multiple trophic levels

Biodiversity has been established as a potential determinant of function in many ecosystems; however, previous research has mostly focused on primary producers and effects at a single trophic level. A broader perspective that considers multiple components of food webs is necessary to understand natural systems. In particular, consumer diversity needs to be more thoroughly examined as trophic interactions and indirect effects can alter ecosystem properties. We test the potential for consumer diversity (fish richness and composition) to govern food web dynamics at two levels of environmental complexity (mesocosms and experimental ponds) and explore the consequences of removing individual species of fish on lower trophic levels. In mesocosms, both the richness and density of zooplankton were reduced when more fish species were present. No effects from the fish treatments were found on phytoplankton, but phosphorus levels increased with higher fish richness. Removing either generalist or specialist fish species increased the richness and density of zooplankton and the amount of phytoplankton, whereas all fish species had redundant effects on nutrients. In ponds, a dominant fish species (specialist shiner) determined the richness and density of zooplankton. In contrast, phytoplankton and nutrients were reduced by higher fish richness in the fall and spring. Overall, the specialist shiner had unique effects on the pond food web suggesting the key to understanding function is the presence of a dominant species and their biological interactions. Differences between mesocosms and ponds are likely due to increased heterogeneity of resources in the ponds allowing species to specialize on different prey. Our study links the biodiversity ecosystem function paradigm with food web concepts to improve predictions for conservation and management actions in response to changes in biodiversity.     

Functional trait diversity across trophic levels determines herbivore impact on plant community biomass

Understanding the consequences of trophic interactions for ecosystem functioning is challenging, as contrasting effects of species and functional diversity can be expected across trophic levels. We experimentally manipulated functional identity and diversity of grassland insect herbivores and tested their impact on plant community biomass. Herbivore resource acquisition traits, i.e. mandible strength and the diversity of mandibular traits, had more important effects on plant biomass than body size. Higher herbivore functional diversity increased overall impact on plant biomass due to feeding niche complementarity. Higher plant functional diversity limited biomass pre‐emption by herbivores. The functional diversity within and across trophic levels therefore regulates the impact of functionally contrasting consumers on primary producers. By experimentally manipulating the functional diversity across trophic levels, our study illustrates how trait‐based approaches constitute a promising way to tackle existing links between trophic interactions and ecosystem functioning.     

Control of arthropod abundance, richness, and composition in a heterogeneous desert city

There is a demand for mechanistic studies to explore underlying drivers behind observed patterns of biodiversity in urban areas. We describe a two-year field experiment in which we manipulated bottom-up (resource availability) and top-down (bird predation) forces on arthropod communities associated with a native plant, Encelia farinosa, across three land-use types—urban, desert remnant, and outlying natural desert—in the Phoenix metropolitan area, Arizona, USA. We monitored the trophic structure, richness, and similarity of the arthropod communities on these manipulated plants over a two-year period. We predicted that (1) increased water resources increase plant productivity, (2) increased productivity increases arthropod abundances, and (3) in the urban habitat, top-down forces are greater than in other habitats and limit arthropod abundances. We also predicted that urban remnant habitats are more similar to urban habitats in terms of arthropod richness and composition. Strong interannual differences due to an unusual cold and dry winter in the first year suppressed plant growth in all but urban habitats, and arthropod abundances in all habitats were severely reduced. In the following year, arthropod abundances in desert and remnant habitats were higher than in urban habitats. Water had positive effects on plant growth and arthropod abundance, but these water effects emerged through complex interactions with habitat type and the presence/absence of cages used to reduce bird predation. Plants grew larger in urban habitats, and phenology also differed between urban and desert habitats. The results from caging suggest that bird predation may not be as important in cities as previously thought, and that arthropods may retard plant growth. As expected, desert communities are strongly bottom-up regulated, but contrary to predictions, we did not find evidence for strong top-down control in the city. Remnant habitats were intermediate between desert and urban habitats in terms of diversity, richness, evenness, arthropod composition and phenology, with urban habitats generally lowest in terms of diversity, richness, and evenness. Our study shows that control of biodiversity is strongly altered in urban areas, influenced by subtle shifts in top-down and bottom-up controls that are often superseded by climatic variations and habitat type.     

Shrubs as ecosystem engineers across an environmental gradient: effects on species richness and exotic plant invasion

Ecosystem-engineering plants modify the physical environment and can increase species diversity and exotic species invasion. At the individual level, the effects of ecosystem engineers on other plants often become more positive in stressful environments. In this study, we investigated whether the community-level effects of ecosystem engineers also become stronger in more stressful environments. Using comparative and experimental approaches, we assessed the ability of a native shrub (Ericameria ericoides) to act as an ecosystem engineer across a stress gradient in a coastal dune in northern California, USA. We found increased coarse organic matter and lower wind speeds within shrub patches. Growth of a dominant invasive grass (Bromus diandrus) was facilitated both by aboveground shrub biomass and by growing in soil taken from shrub patches. Experimental removal of shrubs negatively affected species most associated with shrubs and positively affected species most often found outside of shrubs. Counter to the stress-gradient hypothesis, the effects of shrubs on the physical environment and individual plant growth did not increase across the established stress gradient at this site. At the community level, shrub patches increased beta diversity, and contained greater rarified richness and exotic plant cover than shrub-free patches. Shrub effects on rarified richness increased with environmental stress, but effects on exotic cover and beta diversity did not. Our study provides evidence for the community-level effects of shrubs as ecosystem engineers in this system, but shows that these effects do not necessarily become stronger in more stressful environments.     

Selenium hyperaccumulation reduces plant arthropod loads in the field

The elemental defense hypothesis proposes that some plants hyperaccumulate toxic elements as a defense mechanism. In this study the effectiveness of selenium (Se) as an arthropod deterrent was investigated under field conditions. Arthropod loads were measured over two growing seasons in Se hyperaccumulator habitats in Colorado, USA, comparing Se hyperaccumulator species (Astragalus bisulcatus and Stanleya pinnata) with nonhyperaccumulators (Camelina microcarpa, Astragalus americanus, Descurainia pinnata, Medicago sativa, and Helianthus pumilus). The Se hyperaccumulating plant species, which contained 1000-14 000 microg Se g(-1) DW, harbored significantly fewer arthropods (c. twofold) and fewer arthropod species (c. 1.5-fold) compared with nonhyperaccumulator species that contained < 30 microg Se g(-1) DW. Arthropods collected on Se-hyperaccumulating plants contained three- to 10-fold higher Se concentrations than those found on nonhyperaccumulating species, but > 10-fold lower Se concentrations than their hyperaccumulator hosts. Several arthropod species contained > 100 microg Se g(-1) DW, indicating Se tolerance and perhaps feeding specialization. These results support the elemental defense hypothesis and suggest that invertebrate herbivory may have contributed to the evolution of Se hyperaccumulation.     

Nonnative plant invasion increases urban vegetation structure and influences arthropod communities

Aim

Ecological theory and empirical evidence indicate that greater structural complexity and diversity in plant communities increases arthropod abundance and diversity. Nonnative plants are typically associated with low arthropod abundance and diversity due to lack of evolutionary history. However, nonnative plants increase the structural complexity of forests, as is common in urban forests. Therefore, urban forests are ideal ecosystems to determine whether structural complexity associated with nonnative plants will increase abundance and diversity of arthropods, as predicted by complexity literature, or whether structural complexity associated with nonnative plants will be depauperate of arthropods, as predicted by nonnative plant literature.

Location

We sampled 24 urban temperate deciduous and mixed forests in two cites, Raleigh, North Carolina and Newark, Delaware, in the eastern United States.

Methods

We quantified ground cover vegetation and shrub layer vegetation in each forest and created structural complexity metrics to represent total, nonnative and native understory vegetation structural complexity. We vacuum sampled arthropods from vegetation and quantified the abundance, biomass, richness and diversity of spiders and non-spider arthropods.

Results

Nonnative plants increase understory vegetation complexity in urban forests. In Raleigh and Newark, we found support for the hypotheses that dense vegetation will increase arthropod abundance and biomass, and against the hypothesis that nonnative vegetation will decrease arthropods. Urban forest arthropod abundance and biomass, but not diversity, increased with greater nonnative and native structural complexity.

Main Conclusions

Invaded urban forests may provide adequate food in the form of arthropod biomass to transfer energy to the next trophic level, but likely fail to provide ecological services and functions offered by diverse species, like forest specialists. Urban land managers should survey urban forests for nonnative and native plant communities and prioritize replacing dense nonnative plants with native species when allocating vegetation maintenance resources.     

Effects of bird predation on arthropod abundance and tree growth across an elevational gradient

Considerable uncertainty surrounds the conditions under which birds can cause trophic cascades. In a three‐year experiment, we studied the direct and indirect effects of insectivorous birds on arthropod abundance, herbivory, and growth of striped maple Acer pensylvanicum saplings in a northern hardwood forest of central New Hampshire, USA. We manipulated bird predation by erecting exclosures around saplings and directly manipulated herbivory by removing herbivores. We also examined how climate modifies these interactions by replicating the experiment at three locations along an elevational gradient. Effects of bird predation were variable. Overall, mean arthropod biomass was 20% greater on saplings within bird exclosures than on controls (p<0.05). The mean biomass of leaf‐chewing herbivores, primarily Lepidoptera larvae, was 25% greater within exclosures but not statistically different from controls. To a lesser degree, mean herbivore damage to foliage within exclosures exceeded that of controls but differences were not significant. We also did not detect significant treatment effects on sapling shoot growth. The high understory vegetation density relative to bird abundance, and low rate of herbivory during the study (mean 5% leaf area removed, controls), may have limited the ability of birds to affect sapling growth. Climate effects operated at multiple scales, resulting in a complex interplay of interactions within the food web. Regional synchrony of climatic conditions resulted in annual fluctuations in herbivore abundance and tree growth that were shared across elevations. At the same time, local environmental variation resulted in site differences in the plant, herbivore, and bird communities. These patterns resulted in a mosaic of top–down strengths across time and space, suggesting an overall pattern of limited effects of birds on plant growth, possibly interspersed with hotspots of trophic cascades.