Tree species diversity influences herbivore abundance and damage: meta-analysis of long-term forest experiments

Plant monocultures are commonly believed to be more susceptible to herbivore attacks than stands composed of several plant species. However, few studies have experimentally tested the effects of tree species diversity on herbivory. In this paper, we present a meta-analysis of uniformly collected data on insect herbivore abundance and damage on three tree species (silver birch, black alder and sessile oak) from seven long-term forest diversity experiments in boreal and temperate forest zones. Our aim was to compare the effects of forest diversity on herbivores belonging to different feeding guilds and inhabiting different tree species. At the same time we also examined the variation in herbivore responses due to tree age and sampling period within the season, the effects of experimental design (plot size and planting density) and the stability of herbivore responses over time. Herbivore responses varied significantly both among insect feeding guilds and among host tree species. Among insect feeding guilds, only leaf miner densities were consistently lower and less variable in mixed stands as compared to tree monocultures regardless of the host tree species. The responses of other herbivores to forest diversity depended largely on host tree species. Insect herbivory on birch was significantly lower in mixtures than in birch monocultures, whereas insect herbivory on oak and alder was higher in mixtures than in oak and alder monocultures. The effects of tree species diversity were also more pronounced in older trees, in the earlier part of the season, at larger plots and at lower planting density. Overall our results demonstrate that forest diversity does not generally and uniformly reduce insect herbivory and suggest instead that insect herbivore responses to forest diversity are highly variable and strongly dependent on the host tree species and other stand characteristics as well as on the type of the herbivore.     

Moose and vole browsing patterns in experimentally assembled pure and mixed forest stands

Plants growing in diverse communities are believed to exhibit associational resistance to herbivores, but this hypothesis has seldom been tested experimentally for vertebrate herbivores in forest ecosystems. We examined browsing patterns of the two principal mammalian herbivores of Finnish boreal forests, moose and voles, in young stands where tree species diversity and composition were experimentally manipulated. The stands were composed either of monocultures or different 2–5 species mixtures of Norway spruce, Scots pine, Siberian larch, silver birch, and black alder. Voles and moose showed contrasting responses to stand diversity and species composition. In accordance with the predictions of the associational resistance hypothesis, vole damage was higher in tree monocultures than in mixed stands, although stand diversity effects were statistically significant only at one of the three study areas. Voles also damaged more trees in coniferous than in deciduous stands. In contrast, moose browsing tended to increase with the number of tree species in a stand and with the presence of the preferred tree species, birch, in a mixture. The observed differences in vole and moose responses to stand diversity and species composition are likely to be due to different feeding specialisation, foraging patterns, and movement ability of these herbivores. Voles switched to trees only when the supply of a more preferred food (grasses and forbs) was depleted and restricted their feeding choice only to the most palatable tree species regardless of the number of tree species present per stand. In contrast, tree branches and foliage represented an important part of moose diet throughout the year; moose may be able to tolerate secondary plant metabolites of different tree species better than voles and may thus benefit from diet broadening when more tree species are available. Furthermore, the home range size and foraging ability of these two very differently sized herbivores may partly explain the observed differences in utilisation of different tree species. Finally, both moose and voles showed high spatial and temporal variation in their feeding; in particular, vole damage was more influenced by tree species diversity in areas and years with high vole densities. Thus, diversification of forest stands may have very different effects on mammalian browsing depending on the herbivores present, their densities, and the tree species used in reforestation.     

Wildflower companion plants increase pest parasitation and yield in cabbage fields: Experimental demonstration and call for caution

Monocultures typical of intensive agriculture offer ideal conditions to specialized herbivores while depriving their natural enemies of habitat and nutritional resources. The resulting release of herbivores from both bottom-up and top-down control causes pest outbreaks in economically important crops. Boosting locally occurring natural enemy populations through species-specific habitat management to restore natural herbivore control has been much advocated but remains rarely tested in the field. Here, we investigated whether adding specifically selected flowering plants to monocultures increases parasitation rates of herbivores and crop yield. We performed replicated field experiments in 2 years and found that adding cornflowers (Centaurea cyanus) into cabbage (Brassica oleracea) fields significantly increased larval and egg parasitation and egg predation of the herbivore, reduced herbivory rates, and increased crop biomass in at least 1 year. These findings show that addition of a single, well-chosen flowering plant species can significantly increase natural top-down pest control in monocultures but success is variable. This is relevant on two applied levels. First, well chosen companion plants may partially substitute pesticides in agriculture if the approach is optimized, reducing negative effects such as unspecific killing of non-target organisms, residues in food, contamination of soils and water-bodies and increasing pesticide resistances. Our results suggest that, from an agro-economical point of view, egg parasitoids or predators may be the best targets for habitat management because strong natural selection acts on larval parasitoids to keep their hosts alive for their own development. Second, the addition of non-crop vegetation to monocultures benefits biodiversity conservation directly through resource diversification and indirectly through the reduction of pesticide application that increased natural control makes possible.     

The effects of fire on woody plant encroachment are exacerbated by succession of trees of decreased palatability

The ingression of woody plants into the grassy layer of savannas and grasslands has become a global concern. The increase of woody plants has been primarily attributed over grazing, fire and more recently to the increase of atmospheric CO₂. We used long-term observations and analyses to assess changes in woody vegetation in Ithala Game Reserve (IGR), South Africa. Textural analysis of aerial photographs was used to detect changes in woody vegetation, from 1943 to 2007 in Ithala Game Reserve (IGR), South Africa. Daily rainfall data from 1905 to 2009 were used in a time-series analysis to determine if rainfall patterns have changed. The time-series analysis showed that the low magnitude (0–10 mm) rainfall events decreased from 1916 to 2009 and high magnitude rainfall events increased (10–20 and >20 mm). The mean annual rainfall increased from ～700 to ～850 mm from the 1930s to the 2000s. This change in rainfall was a key factor in the increase in woody vegetation from 1943 to 2009. We also used field data from the same reserve collected over 30 years to assess the increases in tree cover. Tree cover and density increased significantly by 32.5% and 657.9 indiv ha^?1 respectively, over 64 years. Before the proclamation of IGR in 1972, increases in woody vegetation from 1943 were non-significant. After the proclamation of IGR, herbivore population numbers and spatial distribution influenced the accumulation of grassy biomass required to fuel fires. In areas with reduced fuel loads, the consequential suppression of fire accelerated the rate of woody plant invasion into savannas. The increase in woody vegetation coincided with a decrease in palatable (e.g. Acacia gerrardii and Acacia davyi) and an increase in unpalatable woody plants. The avoidance of the unpalatable trees (e.g. Euclea and Searsia species) by large mammalian herbivores has allowed these trees to increase in density relatively unhindered.     

Survival of black alder (Alnus glutinosa L.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) seedlings in a reclaimed oil shale mining area

Early survival and growth of black alder, silver birch and Scots pine were investigated on reclaimed extremely stony and heterogeneous calcareous (pH 8) opencast oil shale mining areas (OOSMAs). Biomass allocation, production, leaf and root adaptations, and mineral nutrition in relation to tree species and soil heterogeneity were analysed. The adaptive strategies of tree species in first-year plantations on OOSMA were different. Scots pine allocated 1.5–2 times more biomass into leaves and fine roots than deciduous trees. The lower leaf/fine root biomass ratio was in proportion to the better survival (%) of seedlings, decreasing in the following order: black alder (93%) ≥ Scots pine (83%) > silver birch (64%). Deciduous trees improved mineral nutrition more by fine-root morphological adaptations than Scots pine; e.g. the mean specific root length (SRL, m g^?1) of short roots increased in the following order: Scots pine (62) < black alder (172) < silver birch (314). The effect of soil heterogeneity on growth and adaptations was minor. All studied species suffered from P and N, and deciduous species also from K deficiency. In the first year after planting, black alder was best adapted to the harsh conditions of the post-mining substrate. The approaches of this study can be used for other regions where wastelands require reclamation.     

Tree species composition rather than diversity triggers associational resistance to the pine processionary moth

The reduction of insect herbivory is one of the services provided by tree diversity in forest ecosystems. While it is increasingly acknowledged that the compositional characteristics of tree species assemblages play a major role in triggering associational resistance to herbivores, underlying mechanisms are less well known. We addressed this question in the ORPHEE experiment by assessing pine processionary moth infestations (Thaumetopoea pityocampa) across a tree diversity gradient from pine monocultures to five species mixtures. We showed that tree species richness per se had no effect on the probability of attack by this pest. By contrast, the infestation rate was strongly dependent on plot composition. Mixtures of pines (Pinus pinaster) and birches (Betula pendula) were less prone to T. pityocampa infestations, whereas mixtures of pines and oaks (Quercus spp.) were more often attacked than pine monocultures. By taking into account the relative height of pines and associated broadleaved species, this effect could be explained by pine apparency. Pines were on average 343 ± 5 cm height. Birches, as fast growing trees, were slightly taller than pines (363 ± 6 cm), while oak trees were significantly smaller (74 ± 1 cm). Host trees of T. pityocampa were then partly hidden in mixtures of pines and birches but more apparent in mixtures with oaks. We suggest that reduced pine apparency disrupted visual cues used by female moths to select host trees prior to oviposition. This study highlights the need to take into account tree traits such as growth rate when selecting the tree species that have to be associated in order to improve forest resistance to pest insects.     

Influence of plant phenostage and ploidy level on oviposition and feeding of two specialist herbivores of spotted knapweed,Centaurea stoebe

A caged field experiment was used to determine how Centaurea stoebe L. phenostage (rosette, single-stem, multiple-stem) and ploidy level (diploid = 2× and tetraploid = 4×) influence oviposition and feeding of two biological control agents, Agapeta zoegana (Lep.: Cochylidae) and Cyphocleonus achates (Col.: Curculionidae). Ploidy level did not influence oviposition patterns of A. zoegana but rosette and one-stem plants had significantly more eggs than multiple-stem (4×) plants. Differences in oviposition levels did not translate into differences in larval densities, but 2× plants (particularly large one-stem plants) had significantly more larvae than 4× plants. There was a significant positive correlation between numbers of larvae and root diameter. Ploidy level and phenostage both had a significant effect on C. achates feeding damage, with adults feeding more frequently on multiple-stem plants. No C. achates larvae were observed when the roots were dissected. Furthermore, the generalist herbivore Arion lusitanicus, naturally present in the garden plots, was predominantly associated with young rosette plants, a stage at which survival rate is acknowledged to be the most important determinant of knapweed density. These results indicate that the combined damage caused by A. zoegana and C. achates, superimposed on damage caused by generalist herbivores in the local community, could provide effective control for C. stoebe.     

Going undercover: increasing canopy cover around a host tree drives associational resistance to an insect pest

Neighbouring heterospecific plants are often observed to reduce the probability of herbivore attack on a given focal plant. While this pattern of associational resistance is frequently reported, experimental evidence for underlying mechanisms is rare particularly for potential plant species diversity effects on focal host plants and their physical environment. Here, we used an established forest diversity experiment to determine whether tree diversity effects on an important insect pest are driven by concomitant changes in host tree growth or the light environment. We examined the effects of tree species richness, canopy cover and tree growth on the probability of occurrence, the abundance, and volume of galls caused by the pineapple gall adelgid Adelges abietis on Norway spruce. Although tree diversity had no effect on gall abundance, we observed that both the probability of gall presence and gall volume (an indicator of maternal fecundity) decreased with tree species richness and canopy cover around host spruce trees. Structural equation models revealed that effects of tree species richness on gall presence and volume were mediated by concurrent increases in canopy cover rather than changes in tree growth or host tree density. As canopy cover did not influence tree or shoot growth, patterns of associational resistance appear to be driven by improved host tree quality or more favourable microclimatic conditions in monocultures compared to mixed‐stands. Our study therefore demonstrates that changes in forest structure may be critical to understanding the responses of herbivores to plant diversity and may underpin associational effects in forest ecosystems.     

Are Tree Species Diversity and Genotypic Diversity Effects on Insect Herbivores Mediated by Ants?

Plant diversity can influence predators and omnivores and such effects may in turn influence herbivores and plants. However, evidence for these ecological feedbacks is rare. We evaluated if the effects of tree species (SD) and genotypic diversity (GD) on the abundance of different guilds of insect herbivores associated with big-leaf mahogany (Swietenia macrophylla) were contingent upon the protective effects of ants tending extra-floral nectaries of this species. This study was conducted within a larger experiment consisting of mahogany monocultures and species polycultures of four species and –within each of these two plot types– mahogany was represented by either one or four maternal families. We selected 24 plots spanning these treatment combinations, 10 mahogany plants/plot, and within each plot experimentally reduced ant abundance on half of the selected plants, and surveyed ant and herbivore abundance. There were positive effects of SD on generalist leaf-chewers and sap-feeders, but for the latter group this effect depended on the ant reduction treatment: SD positively influenced sap-feeders under ambient ant abundance but had no effect when ant abundance was reduced; at the same time, ants had negative effects on sap feeders in monoculture but no effect in polyculture. In contrast, SD did not influence specialist stem-borers or leaf-miners and this effect was not contingent upon ant reduction. Finally, GD did not influence any of the herbivore guilds studied, and such effects did not depend on the ant treatment. Overall, we show that tree species diversity influenced interactions between a focal plant species (mahogany) and ants, and that such effects in turn mediated plant diversity effects on some (sap-feeders) but not all the herbivores guilds studied. Our results suggest that the observed patterns are dependent on the combined effects of herbivore identity, diet breadth, and the source of plant diversity.     

Forage selection by Royle's pika (Ochotona roylei) in the western Himalaya,India

Forage selection decisions of herbivores are often complex and dynamic; they are modulated by multiple cues, such as quality, accessibility and abundance of forage plants. To advance the understanding of plant–herbivore interactions, we explored foraging behavior of the alpine lagomorph Royle's pika (Ochotona roylei) in Kedarnath Wildlife Sanctuary, India. Pika bite counts on food plants were recorded through focal sampling in three permanently marked plots. Food plant abundance was recorded by traditional quadrat procedures; forage selection was estimated with Jacob's selection index. Multiple food-choice experiments were conducted to determine whether forage selection criteria would change with variation in food plant composition. We also analyzed leaf morphology and nutrient content in both major food plants and abundantly available non-food plants. Linear regression models were used to test competing hypotheses in order to identify factors governing forage selection. Royle's pika fed primarily on 17 plant species and each forage selection decision was positively modulated by leaf area and negatively modulated by contents of avoided substances (neutral detergent fiber, acid detergent fiber, acid detergent lignin and tannin) in food plants. Furthermore, significance of the interaction term “leaf size × avoided substance” indicates that plants with large leaves were selected only when they had low avoided substance content. The forage selection criteria did not differ between field and laboratory experiments. The parameter estimates of best fit models indicate that the influence of leaf size or amount of avoided substance on pika forage selection was modulated by the magnitude of predation risk.     

Comparisons of arthropod assemblages on an invasive and native trees: abundance,diversity and damage

The success of exotic plants may be due to lower herbivore loads than those on native plants (Enemies Release Hypothesis). Predictions of this hypothesis include lower herbivore abundances, diversity, and damage on introduced plant species compared to native ones. Greater density or diversity of predators and parasitoids on exotic versus native plants may also reduce regulation of exotic plants by herbivores. To test these predictions, we measured arthropod abundance, arthropod diversity, and foliar damage on invasive Chinese tallow tree (Triadica sebifera) and three native tree species: silver maple (Acer saccharinum), sycamore (Platanus occidentalis), and sweetgum (Liquidambar styraciflua). Arthropod samples were collected with canopy sweep nets from six 20 year old monoculture plots of each species at a southeast Texas site. A total of 2,700 individuals and 285 species of arthropods were caught. Overall, the species richness and abundance of arthropods on tallow tree were similar to the natives. But, ordination (NMS) showed community composition differed on tallow tree compared to all three native trees. It supported an arthropod community that had relatively lower herbivore abundance but relatively more predator species compared to the native species examined. Leaves were collected to determine damage. Tallow tree experienced less mining damage than native trees. The results of this study supported the Enemies Release Hypothesis predictions that tallow tree would have low herbivore loads which may contribute to its invasive success. Moreover, a shift in the arthropod community to fewer herbivores without a reduction in predators may further limit regulation of this exotic species by herbivores in its introduced range.     

Conditional outcomes in plant–herbivore interactions: neighbours matter

Spatial distribution of palatable and unpalatable plants can influence the foraging behaviour of herbivores, thereby changing plant‐damage probabilities. Moreover, the immediate proximity to certain plants can benefit other plants that grow below them, where toxicity or spines act as a physical barrier or concealment against herbivores. This paper presents the results of a multi‐scale experiment performed to test the effect of shrubs as protectors of tree saplings against herbivores and the mechanism involved in Mediterranean ecosystems. We performed a factorial design in two mountain ranges, similar in physiognomy and vegetation, planting saplings of a palatable tree, the maple (Acer opalus subsp. granatense), and an unpalatable tree, the black pine (Pinus nigra), under three different types of shrubs. We considered four experimental microhabitats: highly palatable shrub (Amelanchier ovalis), palatable but spiny shrub (Crataegus monogyna or Prunus ramburii), unpalatable spiny shrub (Berberis vulgaris subsp. australis) and control (gaps of bare soil without shrubs). Three main factors were found to determine the probability of sapling attack: sapling palatability, experimental microhabitat and plot. Palatable saplings (maples) were browsed much more than unpalatable ones (pines). The degree of protection provided by the shrub proved greater as its palatability decreased with respect to sapling palatability, the unpalatable spiny shrub being the safest microhabitat for palatable saplings and bare soil for unpalatable ones. The differences found in number of attacked saplings between plots may be attributable to differences in herbivore pressure. The community context in which interaction takes place, namely the characteristics of the neighbours and the intensity of herbivore pressure, are determining factors for understanding and predicting the damage undergone by a target plant species. The mechanism that best explains these results is associational avoidance of saplings that grow near to unpalatable shrubs. It is necessary to introduce this neighbour effect in theoretical models and food‐web approaches that analyse the plant–herbivore relationships, since it can strongly determine not only the intensity of the interaction, but also the spatial distribution and diversity of the plant community.     

Tree Diversity Limits the Impact of an Invasive Forest Pest

The impact of invasive herbivore species may be lower in more diverse plant communities due to mechanisms of associational resistance. According to the “resource concentration hypothesis” the amount and accessibility of host plants is reduced in diverse plant communities, thus limiting the exploitation of resources by consumers. In addition, the “natural enemy hypothesis” suggests that richer plant assemblages provide natural enemies with more complementary resources and habitats, thus promoting top down regulation of herbivores. We tested these two hypotheses by comparing crown damage by the invasive Asian chestnut gall wasp (Dryocosmus kuriphilus) on chestnut trees (Castanea sativa) in pure and mixed stands in Italy. We estimated the defoliation on 70 chestnut trees in 15 mature stands sampled in the same region along a gradient of tree species richness ranging from one species (chestnut monocultures) to four species (mixtures of chestnut and three broadleaved species). Chestnut defoliation was significantly lower in stands with higher tree diversity. Damage on individual chestnut trees decreased with increasing height of neighboring, heterospecific trees. These results suggest that conservation biological control method based on tree species mixtures might help to reduce the impact of the Asian chestnut gall.     

Habitat selection by endangered Himalayan musk deer (Moschus chrysogaster) and impacts of livestock grazing in Nepal Himalaya: Implications for conservation

Habitat management within and outside protected areas is a key to effective conservation of wildlife. This is particularly vital for declining wildlife populations within the boundary of conservation areas, while sharing their potential habitat range with foraging livestock. In an effort to understand the habitat selection by Himalayan musk deer (Moschus chrysogaster) and explore any potential impacts of livestock grazing, we conducted the present study in a conservation area of central Nepal Himalaya. We recorded data on musk deer and livestock presence and absence (based on signs of fecal pellet, footprint, and resting site) along the elevational transect with associated topographic features (elevation, slope, aspect, distance to water, and vantage point distance) and vegetation features (tree spp., shrub spp., herb spp., and canopy-cover). Using logistic regression model we found that elevation, aspect, canopy-cover, and tree spp. in the area significantly affect the likelihood of habitat selection by musk deer. In particular, they selected the southern aspect of the area with elevation ≥ 3529 m, canopy-cover ≥ 42%, and with stands of Pinus spp. and Abies spp. Slope and canopy-cover significantly affected the foraging area selection by livestock. They selected the gentler slopes in the northern aspect of the area with altitude < 3529 m and canopy-cover < 42%. Also, presence of one group of herbivore (i.e. musk deer and livestock) was not found to affect the likelihood of habitat selection by the other group. These independent habitat selections are possibly the responses to morphological and behavioral adaptations than to impacts and interactions between these two groups of herbivores. We suggest to avoid any disturbances and livestock grazing on the area that disrupt the resources and conditions likely selected and occupied by musk deer population.     

cis-Jasmone indirect action on egg parasitoids (Hymenoptera: Scelionidae) and its application in biological control of soybean stink bugs (Hemiptera: Pentatomidae)

In many plants, the secondary metabolite cis-jasmone activates the metabolic pathway that produces volatile organic compounds attractive to natural enemies and, sometimes, repellent to herbivores. Previous studies indicate that the feeding damage caused by the herbivore Euschistus heros or the exogenous application of cis-jasmone in soybean plants induces the release of herbivore-induced plant volatiles (HIPVs) with a similar chemical profile and these compounds can attract the stink bug egg parasitoid Telenomus podisi (Scelionidae). Herein we tested in field conditions the effect of exogenous application of cis-jasmone in soybean plants on the parasitoid and stink bug community and on stink bug egg parasitism. In two areas, one within a soybean and another within a Crotalaria matrix, we randomly distributed 2 m² plots, with soybean plants induced (treatment, n = 5) or not induced by cis-jasmone (control, n = 5) in the field. We sampled the parasitoid community weekly with yellow sticky traps (n = 3/plot) and monitored parasitism with sentinel eggs of E. heros (n = 150/plot). We also monitored the population of stink bugs weekly, by sampling each plot with shake-cloth technique. The abundance of Scelionidae was highest overall and also in treated plots during the first four weeks in the area with a soybean matrix, but decreased thereafter. The richness of parasitoid families was similar between treatment and control plots in the area with a soybean matrix, but higher in control plots in the area with a Crotalaria matrix. Evenness was higher in control plots in the area with soybean matrix, whereas the reverse occurred in the area with a Crotalaria matrix. Results suggest that treatment with cis-jasmone effectively attracted and enhanced the population of scelionid parasitoids, but had no effect on the occurrence and intensity of parasitism and in the number of stink bugs.     

Effects of stand tree species composition and diversity on abundance of predatory arthropods

Diverse plant communities are predicted to have higher abundance of predators as compared to species-poor ones. In this study we explored whether this prediction holds true for the abundance of predatory arthropods in forest ecosystems, which have been poorly studied in this respect. We collected ground-dwelling arthropods using pitfall traps from six long-term forest diversity experiments in Finland, Sweden and UK. The effects of tree species diversity on abundance of five main groups of predatory arthropods (ants, spiders, carabids, staphylinids and opilionids) were examined by means of meta-analysis. Overall, the diversity of tree species did not affect abundances of predators with the exception of staphylinids, which were more abundant in mixed stands than in monocultures. However, the effects of stand diversity on predator abundance became apparent when analyses were conducted on tree species basis. Preference for stands containing particular tree species was clear in the case of opilionids, carabids and staphylinids, and these preferences overruled the effects of tree species diversity in comparisons between monocultures and mixed stands containing the same tree species. Tree species diversity effects on predator abundance were furthermore mediated by the tree age, plot size and planting density used in the experiment. Overall, our results show that predator abundance does not increase uniformly with increase in tree species diversity, but rather suggest that predators have distinct preferences for stands composed of particular tree species and that these species preferences may overrule the effects of diversity.     

Influence of host plant nitrogen fertilization on hemolymph protein profiles of herbivore Spodoptera exigua and development of its endoparasitoid Cotesia marginiventris

Nitrogen has complex effects on plant–herbivore–parasitoid tritrophic interactions. The negative effects of low nitrogen fertilization in host plants on insect herbivores can be amplified to the higher trophic levels. In the present study, we examined the impact of varying nitrogen fertilization (42, 112, 196, and 280 ppm) of cotton plants (Gossypium hirsutum L.) on the interactions between the beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), and the hymenopteran endoparasitoid Cotesia marginiventris (Cresson) (Hymenoptera: Braconidae). We predicted that the development and fitness of C. marginiventris would be adversely affected by low host plant nitrogen fertilization through the herbivore S. exigua. The percentage of C. marginiventris offspring developing to emerge and spin a cocoon, and total mortality of parasitized S. exigua larvae were unaffected by nitrogen level. The developmental time of C. marginiventris larvae in S. exigua larvae feeding on low (42 ppm) nitrogen cotton plants was approximately 30% longer than that of those feeding on higher (112, 196, and 280 ppm) nitrogen plants. Parasitoid size (length of right metathoracic tibia), a proxy for fitness, of C. marginiventris males was positively affected by nitrogen level. Total amounts of S. exigua hemolymph proteins were not affected by nitrogen level, but were reduced by parasitism by C. marginiventris. Two proteins with molecular weights of ca. 84 and 170 kDa dominated the S. exigua larval hemolymph proteins. Concentrations of the 170 kDa hemolymph protein were unaffected by nitrogen treatment, but parasitism reduced concentrations of the 170 kDa protein. Concentrations of the 84 kDa protein, on the other hand, were interactively affected by parasitism and nitrogen treatment: higher nitrogen fertilization (112, 196, and 280 ppm) increased protein concentrations relative to the 42 ppm treatment for unparasitized S. exigua larvae, whereas nitrogen treatment had no effects on parasitized larvae. For S. exigua larvae feeding on 42 ppm nitrogen plants, parasitism increased concentration of the 84 kDa protein, while for those feeding on 112, 196, and 280 ppm nitrogen plants, parasitism decreased concentrations of the protein. Possible mechanisms and ecological consequences for the extended development of C. marginiventris on S. exigua hosts grown on low-nitrogen plants are discussed.     

Flower-heads, herbivores, and their parasitoids: food web structure along a fertility gradient

Abstract. 1. The ways in which a soil fertility gradient affects three trophic level food webs defined by plants of the family Asteraceae, flower‐head herbivores, and their parasitoids was investigated. It was tested how the fertility gradient alters: (i) the abundance and richness of plants, herbivores, and their parasitoids, (ii) the herbivore–plant ratio, and (iii) the connectance of the plant–herbivore community matrix. 2. From April to May 2000, plants and insects were sampled in 16 Brazilian Cerrado (sensu stricto) sites along a physiognomic gradient varying from open shrublands (cerrado) to closed woodlands (cerradão). Sites were objectively positioned along the physiognomic gradient by a single index, tree density. Sixty‐seven per cent of the variation in tree density among sites was correlated to two principal components of a PCA, representing gradients of soil fertility. 3. Asteraceae abundance, richness, and flower‐head availability were negatively related to tree density due to their preference for sunny environments, despite the surplus of soil nutrients. The abundance and richness of Diptera and Lepidoptera, the most important flower‐head herbivores, were also negatively related to tree density. Parasitoid abundance decreased with tree density; however, the number of parasitoids per hosts was lower in cerrado, suggesting that top‐down forces are not getting stronger in more productive sites, as could be expected. 4. Community allometry analyses showed that the herbivore to plant ratio was independent of community richness and did not respond to tree density. 5. Connectance of the plant–herbivore matrix was dependent on the community matrix size. Proportionally, species‐rich cerrado sites had fewer interactions than their species‐poor counterparts. Nevertheless, after removing the effect of the matrix size, connectance was not related to tree density. 6. Soil fertility, as the primary cause of the cerrado–cerradão physiognomic gradient, strongly affected the abundance and richness of plants, herbivores and their parasitoids; however, it had little effect on important community attributes, such as the herbivore–plant ratio and the connectance of the plant–herbivore matrix.     

Termite mound cover and abundance respond to herbivore‐mediated biotic changes in a Kenyan savanna

Both termites and large mammalian herbivores (LMH) are savanna ecosystem engineers that have profound impacts on ecosystem structure and function. Both of these savanna engineers modulate many common and shared dietary resources such as woody and herbaceous plant biomass, yet few studies have addressed how they impact one another. In particular, it is unclear how herbivores may influence the abundance of long‐lived termite mounds via changes in termite dietary resources such as woody and herbaceous biomass. While it has long been assumed that abundance and areal cover of termite mounds in the landscape remain relatively stable, most data are observational, and few experiments have tested how termite mound patterns may respond to biotic factors such as changes in large herbivore communities. Here, we use a broad tree density gradient and two landscape‐scale experimental manipulations—the first a multi‐guild large herbivore exclosure experiment (20 years after establishment) and the second a tree removal experiment (8 years after establishment)—to demonstrate that patterns in Odontotermes termite mound abundance and cover are unexpectedly dynamic. Termite mound abundance, but areal cover not significantly, is positively associated with experimentally controlled presence of cattle, but not wild mesoherbivores (15–1,000 kg) or megaherbivores (elephants and giraffes). Herbaceous productivity and tree density, termite dietary resources that are significantly affected by different LMH treatments, are both positive predictors of termite mound abundance. Experimental reductions of tree densities are associated with lower abundances of termite mounds. These results reveal a richly interacting web of relationships among multiple savanna ecosystem engineers and suggest that termite mound abundance and areal cover are intimately tied to herbivore‐driven resource availability.