Ranking Lepidopteran Use of Native Versus Introduced Plants

Abstract:  In light of the wide-scale replacement of native plants in North America with introduced, invasive species and noninvasive ornamental plants that evolved elsewhere, we compared the value of native and introduced plants in terms of their ability to serve as host plants for Lepidoptera. Insect herbivores such as Lepidoptera larvae are critically important components of terrestrial food webs and any reduction in their biomass or diversity due to the loss of acceptable host plants is predicted to reduce the production of the many insectivores in higher trophic levels. We conducted an exhaustive search of host records in the literature. We used the data we gathered to rank all 1385 plant genera that occur in the mid-Atlantic states of the United States by their ability to support Lepidoptera richness. Statistical comparisons were made with Welch's test for equality of means. Woody plants supported more species of moths and butterflies than herbaceous plants, native plants supported more species than introduced plants, and native woody plants with ornamental value supported more Lepidoptera species than introduced woody ornamentals. All these differences were highly significant. Our rankings provide a relative measure that will be useful for restoration ecologists, landscape architects and designers, land managers, and landowners who wish to raise the carrying capacity of particular areas by selecting plants with the greatest capacity for supporting biodiversity.     

Impacts of Alternative Timber Harvest Practices on Leaf-Chewing Herbivores of Oak

Abstract:  Studies of the effects of logging on Lepidoptera rarely address landscape-level effects or effects on larval, leaf-feeding stages. We examined the impacts of uneven-aged and even-aged logging on the abundance, richness, and community structure of leaf-chewing insects of white ( Quercus alba L.) and black ( Q. velutina L.) oak trees remaining in unharvested areas by sampling 3 years before and 7 years after harvest. After harvest, white oaks in uneven-aged sites had 32% fewer species of leaf-chewing insects than control sites. This reduction in species richness may have resulted from changes in microclimate (reducing plant quality and/or changing leaf phenology) that affected a much larger total area of each site than did even-aged cuts. For black oak after harvest, species richness in uneven- and even-aged sites increased relative to levels before harvest. Harvesting did not alter total insect density or community structure in the unlogged habitat for either oak species with one exception: insect density on black oak increased in the oldest forest block. Community structure of herbivores of black and white oaks in clearcut gaps differed from that of oaks in intact areas of even-aged sites. Furthermore, both richness and total insect density of black oaks were reduced in clearcut gaps. We suggest that low-level harvests alter herbivore species richness at the landscape level. Treatment effects were subtle because we sampled untreated areas of logged landscapes, only one harvest had occurred, and large temporal and spatial variation in abundance and richness existed. Although the effects of logging were greater in uneven-aged sites, the effects of even-aged management are likely to increase as harvesting continues.     

青岛市城市绿地种子植物区系的构成特征

在资料分析和2013年夏秋二季进行的实地调查基础上研究了青岛市城市绿地种子植物区系的科属种构成、种的生活型构成、属的地理分布区类型构成,并比较了青岛市城市绿地种子植物区系与青岛崂山种子植物区系、中国种子植物区系属的地理分布区类型构成差异。结果表明,青岛市城市绿地种子植物区系有种子植物69科155属228种,有含5种及以上的12个较大科、124个单种属、183种木本植物和45种草本植物,有热带分布属32属、温带分布属95属、世界分布属14属和中国特有属14属。青岛市城市绿地种子植物区系有植物种类丰富多样,木本植物种类多,区系中属、种的分化程度高,属的地理分布区类型多样且具有很典型的温带特征和鲜明的中国特色等特点。     

Herbivores on a dominant understory shrub increase local plant diversity in rain forest communities

Indirect effects of trophic interactions on biodiversity can be large and common, even in complex communities. Previous experiments with dominant understory Piper shrubs in a Costa Rican rain forest revealed that increases in herbivore densities on these shrubs caused widespread seedling mortality as a result of herbivores moving from Piper to seedlings of many different plant genera. We tested components of the Janzen-Connell hypothesis by conducting focused studies on the effects of specialist and generalist Piper herbivores on local seedling diversity. Whereas specialist herbivores are predicted to increase mortality to neighboring seedlings that are closely related to the source plant, true generalists moving from source plants may cause density-dependent mortality of many species, and possibly increase richness if new species replace abundant species that have been thinned by herbivores. Therefore, we hypothesized that seedling richness would be greater in understory control plots created in patches of Piper that had normal densities of generalist herbivores compared to plots from which we removed generalist herbivores manually from all Piper shrubs. After 15 months, generalist-herbivore-removal plots had > 40% fewer seedlings, > 40% fewer species, and 40% greater seedling evenness, on average, than control plots with generalist herbivores intact. Using a complementary approach in unmanipulated plots in four forests, we used path analysis to test for a positive association between seedling diversity and herbivore damage on Piper species. In unmanipulated plots, for both generalist and specialist herbivores, our data were significant fits to the causal model that Piper herbivores decrease evenness and increase plant species richness, corroborating the experimental results. Because herbivores changed how individuals were apportioned among the species and families present (lower evenness), one interpretation of these associations between herbivores on Piper shrubs and local seedling richness is that high seedling mortality in dominant families allowed the colonization or survival of less common species. If interspecific or apparent competition allowed for a relative increase in species richness, then the Janzen-Connell hypothesis may extend its predictions to generalist seedling predators. We speculate that apparent competition may explain some of the deviations from neutral model predictions, especially at small scales.     

Bottom-up multitrophic effects in resprouting plants

Severe damage often provokes compensatory resprouting of plants, which commonly modify plant morphological and phenological traits. Rapid plant growth often results in poorly defended nutrient-rich foliage, which is more susceptible to foliar-chewing herbivores. It is less known how other guilds of arthropods are affected by plant regrowth. We tested the hypotheses that clipping-induced resprouting and nutrient availability, separately and in combination, would (1) influence plant traits, (2) benefit chewing herbivores, sap-suckers, gallers, and pre-dispersal seed predators, and (3) cascade up to the third trophic level by positively affecting herbivores. Resprouted plants were morphologically and phenologically different from undamaged plants; as a result, seed predation, infestation rate, richness, and diversity of seed predators increased, and species composition was altered. Leaf consumption by chewing herbivores was four times higher on resprouted plants. The number of galls decreased, whereas the abundance of sap-sucking and leaf-chewing insects was not affected. The incidence of predators and parasitoids was also higher on resprouted plants and on plants with nutrients added, but the increase was less pronounced compared to the herbivores they feed on. Thus, the effects of resprouting, contingent on nutrient availability, can propagate simultaneously through two independent tri-trophic level pathways.     

Differential responses of vertebrate and invertebrate herbivores to traits of New Zealand subalpine shrubs

Plant traits are influenced by herbivore diet selection, but little is known about how traits are affected by different types of herbivores. We related eight traits of 27 subalpine shrub species in South Island, New Zealand, to damage of these shrubs by introduced red deer (Cervus elaphus) and native invertebrate herbivores using phylogenetically explicit modeling. Deer preferentially consumed species that grew quickly, were low in foliar tannins, or had high leaf area per unit mass. However, these traits did not trade off against each other; rather, they could be related to different multivariate defense strategies. Although the proportion of leaves damaged by leaf-chewing invertebrates also increased with stem growth, invertebrates did not damage the same fast growing species as those preferred by deer. Other traits may also be important in determining herbivore preferences, as suggested by the high proportion of variation in herbivory explained by phylogeny. Last, we found that the composition of invertebrate herbivore communities was more similar among closely related shrubs, and consequently, the range of invertebrate-plant associations may change if introduced deer shift plant composition toward slow-growing species. Overall, our results demonstrate the importance of herbivore type and coevolved interactions for the adaptive significance of plant traits.     

Direct and indirect effects of CO2, nitrogen, and community diversity on plant-enemy interactions

Resource abundance and plant diversity are two predominant factors hypothesized to influence the amount of damage plants receive from natural enemies. Many impacts of these environmental variables on plant damage are likely indirect and result because both resource availability and diversity can influence plant traits associated with attractiveness to herbivores or susceptibility to pathogens. We used a long-term, manipulative field experiment to investigate how carbon dioxide (CO2) enrichment, nitrogen (N) fertilization, and plant community diversity affect plant traits and the amount of herbivore and pathogen damage experienced by the common prairie legume Lespedeza capitata. We detected little evidence that CO2 or N affected plant traits; however, plants growing in high-diversity treatments (polycultures) were taller, were less pubescent, and produced thinner leaves (higher specific leaf area). Interestingly, we also detected little evidence that CO2 or N affect damage. Plants growing in polycultures compared to monocultures, however, experienced a fivefold increase in damage from generalist herbivores, 64% less damage from specialist herbivores, and 91% less damage from pathogens. Moreover, within diversity treatments, damage by generalist herbivores was negatively correlated with pubescence and often was positively correlated with plant height, while damage by specialist herbivores typically was positively correlated with pubescence and negatively associated with height. These patterns are consistent with changes in plant traits driving differences in herbivory between diversity treatments. In contrast, changes in measured plant traits did not explain the difference in disease incidence between monocultures and polycultures. In summary, our data provide little evidence that CO2 or N supply alter damage from natural enemies. By contrast, plants grown in monocultures experienced greater specialist herbivore and pathogen damage but less generalist herbivore damage than plants grown in diverse communities. Part of this diversity effect was mediated by changes in plant traits, many of which likely are plastic responses to diversity treatments, but some of which may be the result of evolutionary changes in response to these long-term experimental manipulations.     

Plant chemistry and insect sequestration

Most plant families are distinguished by characteristic secondary metabolites, which can function as putative defence against herbivores. However, many herbivorous insects of different orders can make use of these plant-synthesised compounds by ingesting and storing them in their body tissue or integument. Such sequestration of putatively unpalatable or toxic metabolites can enhance the insects’ own defence against enemies and may also be involved in reproductive behaviour. This review gives a comprehensive overview of all groups of secondary plant metabolites for which sequestration by insect herbivores belonging to different orders has been demonstrated. Sequestered compounds include various aromatic compounds, nitrogen-containing metabolites such as alkaloids, cyanogenic glycosides, glucosinolates and other sulphur-containing metabolites, and isoprenoids such as cardiac glycosides, cucurbitacins, iridoid glycosides and others. Sequestration of plant compounds has been investigated most in insects feeding or gathering on Apocynaceae s.l. (Apocynoideae, Asclepiaoideae), Aristolochiaceae, Asteraceae, Boraginaceae, Fabaceae and Plantaginaceae, but it also occurs for some gymnosperms and even lichens. In total, more than 250 insect species have been shown to sequester plant metabolites from at least 40 plant families. Sequestration predominates in the Coleoptera and Lepidoptera, but also occurs frequently in the orders Heteroptera, Hymenoptera, Orthoptera and Sternorrhyncha. Patterns of sequestration mechanisms for various compound classes and common or individual features occurring in different insect orders are highlighted. More research is needed to elucidate the specific transport mechanisms and the physiological processes of sequestration in various insect species.     

Effects of Growth Form and Functional Traits on Response of Woody Plants to Clearing and Fragmentation of Subtropical Rainforest

The conservation implications of large‐scale rainforest clearing and fragmentation on the persistence of functional and taxonomic diversity remain poorly understood. If traits represent adaptive strategies of plant species to particular circumstances, the expectation is that the effect of forest clearing and fragmentation will be affected by species functional traits, particularly those related to dispersal. We used species occurrence data for woody plants in 46 rainforest patches across 75,000 ha largely cleared of forest by the early 1900s to determine the combined effects of area reduction, fragmentation, and patch size on the taxonomic structure and functional diversity of subtropical rainforest. We compiled species trait values for leaf area, seed dry mass, wood density, and maximum height and calculated species niche breadths. Taxonomic structure, trait values (means, ranges), and the functional diversity of assemblages of climbing and free‐standing plants in remnant patches were quantified. Larger rainforest patches had higher species richness. Species in smaller patches were taxonomically less related than species in larger patches. Free‐standing plants had a high percentage of frugivore dispersed seeds; climbers had a high proportion of small wind‐dispersed seeds. Connections between the patchy spatial distribution of free‐standing species, larger seed sizes, and dispersal syndrome were weak. Assemblages of free‐standing plants in patches showed more taxonomic and spatial structuring than climbing plants. Smaller isolated patches retained relatively high functional diversity and similar taxonomic structure to larger tracts of forest despite lower species richness. The response of woody plants to clearing and fragmentation of subtropical rainforest differed between climbers and slow‐growing mature‐phase forest trees but not between climbers and pioneer trees. Quantifying taxonomic structure and functional diversity provides an improved basis for conservation planning and management by elucidating the effects of forest‐area reduction and fragmentation. Efectos de la Forma de Crecimiento y Atributos Funcionales en la Respuesta de Plantas Leñosas al Desmonte y Fragmentación de Bosque Lluvioso Subtropical     

Forest age influences oak insect herbivore community structure, richness, and density.

Plant succession is one of many factors that may affect the composition and structure of herbivorous insect communities. However, few studies have examined the effect of forest age on the diversity and abundance of insect communities. If forest age influences insect diversity, then the schedule of timber harvest rotation may have consequent effects on biodiversity. The insect herbivore community on Quercus alba (white oak) in the Missouri Ozarks was sampled in a chronoseries, from recently harvested (2 yr) to old-growth (approximately 313 yr) forests. A total of nine sites and 39 stands within those sites were sampled in May and August 2003. Unique communities of plants and insects were found in the oldest forests (122-313 yr). Density and species richness of herbivores were positively correlated with increasing forest age in August but not in May. August insect density was negatively correlated with heat load index; in addition, insect density and richness increased over the chronoseries, but not on the sunniest slopes. Forest structural diversity (number of size classes) was positively correlated with forest age, but woody plant species richness was not. In sum, richness, density, and community structure of white oak insect herbivores are influenced by variation in forest age, forest structure, relative abundance of plant species, and abiotic conditions. These results suggest that time between harvests of large, long-lived, tree species such as white oak should be longer than current practice in order to maintain insect community diversity.     

Species-specific responses of herbivores to within-plant and environmentally mediated between-plant variability in plant chemistry

Allocation of resources to growth and defense against herbivores crucially affects plant competitiveness and survival, resulting in a specific distribution of assimilates and defense compounds within plant individuals. Additionally, plants rarely experience stable environmental conditions, and adaptations to abiotic and biotic stresses may involve shifts in resistance to herbivores. We studied the allocation of phytochemicals in Brassica oleracea (Brussels sprouts) due to leaf age, drought stress and herbivore damage and assessed effects on two lepidopteran herbivores differing in diet breadth: the generalist Spodoptera littoralis and the specialist Pieris brassicae. Glucosinolates as secondary defense compounds and total nitrogen and carbon were quantified and linked to plant palatability, i.e., herbivore feeding preference. Herbivore responses were highly species-specific and partially related to changes in phytochemicals. Spodoptera littoralis preferred middle-aged leaves with intermediate levels of glucosinolates and nitrogen over young, glucosinolate and nitrogen rich leaves, as well as over old leaves, poor in glucosinolates and nitrogen. In contrast, P. brassicae preferred young leaves. Both species preferred severely drought-stressed plants to the well-watered control, although analyzed glucosinolate concentrations did not differ. Both S. littoralis and P. brassicae feeding induced an increase of indole glucosinolate levels, which may explain a reduced consumption of damaged plants detected for S. littoralis but not for P. brassicae. By revealing distinct, sometimes contrasting responses of two insect herbivores to within-plant and stress-mediated intraspecific variation in phytochemistry of B. oleracea, this study emphasizes the need to consider specific herbivore responses to understand and predict the interactions between herbivores and variable plants.     

Leaf quality, predators, and stochastic processes in the assembly of a diverse herbivore community

Ecological communities are structured by both deterministic, niche-based processes and stochastic processes such as dispersal. A pressing issue in ecology is to determine when and for which organisms each of these types of processes is important in community assembly. The roles of deterministic and stochastic processes have been studied for a variety of communities, but very few researchers have addressed their contribution to insect herbivore community structure. Insect herbivore niches are often described as largely shaped by the antagonistic pressures of predation and host plant defenses. However host plants are frequently discrete patches of habitat, and their spatial arrangement can affect herbivore dispersal patterns. We studied the roles of predation, host plant quality, and host spatial proximity for the assembly of a diverse insect herbivore community on Quercus alba (white oak) across two growing seasons. We examined abundances of feeding guilds to determine if ecologically similar species responded similarly to variation in niches. Most guilds responded similarly to leaf quality, preferring high-nitrogen, low-tannin host plants, particularly late in the growing season, while bird predation had little impact on herbivore abundance. The communities on the high-quality plants tended to be larger and, in some cases, have greater species richness. We analyzed community composition by correlating indices of community similarity with predator presence, leaf quality similarity, and host plant proximity. Birds did not affect community composition. Community similarity was significantly associated with distance between host plants and uncorrelated with leaf quality similarity. Thus although leaf quality significantly affected the total abundance of herbivores on a host plant, in some cases leading to increased species richness, dispersal limitation may weaken this relationship. The species composition of these communities may be driven by stochastic processes rather than variation in host plant characteristics or differential predation by insectivorous birds.     

Are lower-latitude plants better defended? Palatability of freshwater macrophytes

Increased herbivory at lower latitudes is hypothesized to select for more effective plant defenses. Feeding assays with seaweeds and salt marsh plants support this hypothesis, with low-latitude plants experiencing greater damage in the field and being less palatable than higher-latitude plants. We tested this hypothesis for freshwater macrophytes because they offered an independent plant lineage and habitat type for testing this general hypothesis and because the patchiness of consumer occupancy across isolated water bodies might produce local variance in herbivory that would override geographic variance and produce different results for this habitat type. When we fed eight congeneric pairs of live plants from four sites in Indiana vs. four sites in South Florida (-215 and 0 frost days/yr respectively) to three species of crayfishes and one species of snail, three of the four herbivores significantly preferred high-latitude to low-latitude plants. For two crayfishes that differed in feeding on live plants (one favoring high-latitude plants and one not), we retested feeding using foods composed of freeze-dried and finely ground plants, thus removing structural characteristics while retaining most chemical/nutritional traits. In this assay, both herbivores strongly preferred high-latitude plants, suggesting that lower-latitude plants had been selected for more deterrent chemical traits. When we collected 22 pairs of congeneric plants from 9 sites throughout Indiana vs. 13 sites in Central Florida (-215 and -95 frost days/yr respectively) and tested these in feeding assays with three crayfishes using dried, ground, and reconstituted plant material, we found a significant effect of latitude for only one of three species of herbivore. Overall, our results suggest a preference for high-latitude plants, but the strength of this relationship varied considerably across small scales of latitude that differed considerably in numbers of frost-free days. The difference in results suggests that large changes in frost frequency over small spatial scales may affect selection for plant defenses, that local variance in herbivory overrode differential selection at geographic scales, or that these possibilities interact when durations of cold weather periodically exclude herbivores from shallower habitats, producing heterogeneous selection for defenses at small spatial scales.     

云南地区中国种子植物特有属的研究

物种特有现象的研究具有重要的区系地理学和保护生物学意义。利用大尺度的植物物种分布信息,探讨了云南地区中国种子植物特有属的科、属组成,区系性质及其地理分布格局。结果表明,云南地区拥有中国种子植物特有属125属,分属于59科,共含种子植物246种,是中国种子植物特有属的重要分布地区之一。含特有属较多的科主要为菊科Compositae、苦苣苔科Gesnefiaceae、伞形科Umbelliferae、禾本科Gramineae和唇形科Labiatae。从各属所包含的物种数来看,单型属占主体地位。根据科的分布区类型构成,云南地区中国种子植物特有属主要起源于热带地区。从属的生活型构成来看,主要以草本植物属为主,而木本植物属所占比重相对较小,暗示着研究区域内特有属的区系兼有年轻成分和古老成分,但以年轻成分为主。从空间分布格局来看,特有属主要分布在“田中线”以东地区。与过去的研究结果不同,我们发现云南地区拥有3个重要的特有属分布中心和3个次级分布中心。     

井冈山猴头杜鹃群落特征的研究

以井冈山猴头杜鹃（Rhododendron simiarum）群落5个样地调查资料为基础,对该群落的物种组成、区系成分、群落结构和物种多样性进行了研究,结果表明：群落有维管植物30科43属50种,其中蕨类植物有5科6属6种,裸子植物4科4属4种,被子植物21科33属40种。区系分析显示该植物区系起源古老,地理成分复杂,以热带性成分为主,温带性成分占有一定的比例。群落为复层异龄结构,猴头杜鹃、甜槠（Castanopsis eyrei）、福建柏（Fokienia hodginsii）、台湾松（Pinus taiwanensis）、薯豆（Elaeocarpus japonicus）和白豆杉（Pseudotaxus chienii）为群落的建群种。除Pielou指数外,群落物种多样性指数较低,各样地间多样性指数测值变幅较大;测度猴头杜鹃群落物种多样性时,各种多样性指数基本表现出相同的趋势;总体来看,猴头杜鹃群落物种多样性在群落不同层次的变化规律为：灌木层〉乔木层〉草本层。     

Predator diversity and identity drive interaction strength and trophic cascades in a food web

Declining predator diversity may drastically affect the biomass and productivity of herbivores and plants. Understanding how changes in predator diversity can propagate through food webs to alter ecosystem function is one of the most challenging ecological research topics today. We studied the effects of predator removal in a simple natural food web in the Sierra Nevada mountains of California (USA). By excluding the predators of the third trophic level of a food web in a full-factorial design, we monitored cascading effects of varying predator diversity and composition on the herbivorous beetle Chrysomela aeneicollis and the willow Salix orestera, which compose the first and second trophic levels of the food web. Decreasing predator diversity increased herbivore biomass and survivorship, and consequently increased the amount of plant biomass consumed via a trophic cascade. Despite this simple linear mean effect of diversity on the strength of the trophic cascade, we found additivity, compensation, and interference in the effects of multiple predators on herbivores and plants. Herbivore survivorship and predator-prey interaction strengths varied with predator diversity, predator identity, and the identity of coexisting predators. Additive effects of predators on herbivores and plants may have been driven by temporal niche separation, whereas compensatory effects and interference occurred among predators with a similar phenology. Together, these results suggest that while the general trends of diversity effects may appear linear and additive, other information about species identity was required to predict the effects of removing individual predators. In a community that is not temporally well-mixed, predator traits such as phenology may help predict impacts of species loss on other species. Information about predator natural history and food web structure may help explain variation in predator diversity effects on trophic cascades and ecosystem function.     

Screening for induced herbivore resistance in Swedish intertidal seaweeds

Terrestrial plants have long been known to induce resistance towards herbivores in response to direct grazing, and strong evidence of inter-plant information transfer through volatile signals has been reported recently. Still, little is known about information exchange in aquatic plant–herbivore interactions. In this study, 12 Swedish seaweed species were exposed either to direct grazing by a generalist crustacean herbivore (Idotea granulosa), or to waterborne signals produced by actively feeding herbivores for 1 week. In order to test for the presence of induced chemical resistance in the different seaweed species, the dried and homogenized seaweed tissues were incorporated into an agar matrix, and herbivores were allowed to choose between the resulting control and induced artificial diets in two different two-choice feeding trials. The herbivores were actively feeding from all seaweed species in the induction experiments, although the amount of seaweed tissue consumed differed significantly between species. A chemically based induced herbivore resistance was found in response to direct grazing in four of the tested seaweed species (two red, one brown, and one green seaweed species). Furthermore, four seaweeds (one red, two brown, and one green seaweed species) induced resistance towards further grazing in response to waterborne chemical signals. Several seaweed species responded differently when exposed to different herbivore-related cues, indicating that both cues and responses can be highly specific. The results show that herbivore-induced resistance is present in 7 of 12 of the tested Swedish seaweed species, but that different signals (i.e., direct grazing and waterborne cues) elicit complex responses in the seaweeds.     

Rainfall and soils modify plant community response to grazing in Serengeti National Park

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.     

The Silent Mass Extinction of Insect Herbivores in Biodiversity Hotspots

Abstract: Habitat loss is silently leading numerous insects to extinction. Conservation efforts, however, have not been designed specifically to protect these organisms, despite their ecological and evolutionary significance. On the basis of species–host area equations, parameterized with data from the literature and interviews with botanical experts, I estimated the number of specialized plant‐feeding insects (i.e., monophages) that live in 34 biodiversity hotspots and the number committed to extinction because of habitat loss. I estimated that 795,971–1,602,423 monophagous insect species live in biodiversity hotspots on 150,371 endemic plant species, which is 5.3–10.6 monophages per plant species. I calculated that 213,830–547,500 monophagous species are committed to extinction in biodiversity hotspots because of reduction of the geographic range size of their endemic hosts. I provided rankings of biodiversity hotspots on the basis of estimated richness of monophagous insects and on estimated number of extinctions of monophagous species. Extinction rates were predicted to be higher in biodiversity hotspots located along strong environmental gradients and on archipelagos, where high spatial turnover of monophagous species along the geographic distribution of their endemic plants is likely. The results strongly support the overall strategy of selecting priority conservation areas worldwide primarily on the basis of richness of endemic plants. To face the global decline of insect herbivores, one must expand the coverage of the network of protected areas and improve the richness of native plants on private lands.     

Asymmetric competition via induced resistance: specialist herbivores indirectly suppress generalist preference and populations

Species may compete indirectly by altering the traits of a shared resource. For example, herbivore-induced responses in plants may make plants more resistant or susceptible to additional herbivorous insect species. Herbivore-induced plant responses can significantly affect interspecific competition and herbivore population dynamics. These herbivore-herbivore indirect interactions have been overlooked in aquatic ecosystems where previous studies used the same herbivore species to induce changes and to assess the effects of these changes. We asked whether seaweed grazing by one of two herbivorous, congeneric snail species (Littorina obtusata or Littorina littorea) with different feeding strategies and preferences would affect subsequent feeding preferences of three herbivore species (both snails and the isopod Idotea baltica) and population densities of three herbivore species (both snails and a third periwinkle snail, Lacuna vincta). In addition, we measured phlorotannin concentrations to test the hypothesis that these metabolites function as induced defenses in the Phaeophyceae. Snail herbivory induced cue-specific responses in apical tissues of the seaweed Fucus vesiculosus that affected the three herbivore species similarly. When compared to ungrazed controls, direct grazing by Littorina obtusata reduced seaweed palatability by at least 52% for both snail species and the isopod species. In contrast, direct grazing by L. littorea did not decrease seaweed palatability for any herbivore, indicating herbivore-specific responses. Previous grazing by L. obtusata reduced populations of L. littorea on outplanted seaweeds by 46% but had no effect on L. obtusata populations. Phlorotannins, a potential class of inducible chemicals in brown algae, were not more concentrated in grazed seaweed tissues, suggesting that some other trait was responsible for the induced resistance. Our results indicate that marine herbivores may compete via inducible responses in shared seaweeds. These plant-mediated interactions were asymmetric with a specialist (L. obtusata) competitively superior to a generalist (L. littorea).