The effects of herbivory on neighbor interactions along a coastal marsh gradient

Many current theories of community function are based on the assumption that disturbances such as herbivory act to reduce the importance of neighbor interactions among plants. In this study, we examined the effects of herbivory (primarily by nutria, Myocastor coypus) on neighbor interactions between three dominant grasses in three coastal marsh communities, fresh, oligohaline, and mesohaline. The grasses studied were Panicum virgatum, Spartina patens, and Spartina alterniflora, which are dominant species in the fresh, oligohaline, and mesohaline marshes, respectively. Additive mixtures and monocultures of transplants were used in conjunction with exclosure fences to determine the impact of herbivory on neighbor interactions in the different marsh types. Herbivory had a strong effect on all three species and was important in all three marshes. In the absence of herbivores, the impact of neighbors was significant for two of the species (Panicum virgatum and Spartina patens) and varied considerably between environments, with competition intensifying for Panicum virgatum and decreasing for Spartina patens with increasing salinity. Indications of positive neighbor effects (mutualisms) were observed for both of these species, though in contrasting habitats and to differing degrees. In the presence of herbivores, however, competitive and positive effects were eliminated. Overall, then, it was observed that in this case, intense herbivory was able to override other biotic interactions such as competition and mutualism, which were not detectable in the presence of herbivores.     

Leaf nutrient content and host plant selection by leaf-cutter ants, Atta laevigata, in a Neotropical savanna

The selection of nutrient-rich leaves by leaf-cutter ants ( Atta spp.) is thought to be of indirect benefit to these ants by promoting the growth of their symbiotic fungus. However, relatively few studies have analyzed the influence of leaf nutrient content on host plant selection by leaf-cutter ants, and conflicting results have been found. We compared the content of eight nutritional elements plus the non-nutrient aluminum between leaves harvested by colonies of Atta laevigata (Smith) (Hymenoptera: Formicidae: Attini) and leaves collected randomly within their foraging areas. In addition, we evaluated whether leaf nutrient content explained the frequency with which these ants attacked and defoliated some of the tree species found in the study area. For 2 years, we monitored 17–26 trees from 15 species and determined the number of times each plant was attacked and the amount and type of foliage removed. Leaves harvested by A. laevigata presented significantly higher concentrations of N, P, K, Zn, and Cu than those collected randomly. This result is likely to reflect the foraging pattern presented by these ants, which were selective both in terms of the plant species and age of leaves most commonly attacked. Young leaves were the only or the main leaf type exploited in many species, and in comparison to mature leaves these presented significantly higher concentrations of P and K. Large differences in the mean number of ant attacks on the tree species studied were also observed, and those presenting more leaf N tended to be the most frequently attacked.     

The importance of plant relatedness for host utilization among phytophagous insects

Current methods for measuring similarity among phytophagous insect communities fail to consider the phylogenetic relationship between host plants. We analysed this relation based on 3580 host observations of 1174 beetle species associated with 100 species of angiosperms in two different forest types in Panama. We quantified the significance of genetic distance as well as taxonomic rank among angiosperms in relation to species overlap in beetle assemblages. A logarithmic model describing the decrease in beetle species similarity between host-plant species of increasing phylogenetic distance explains 35% of the variation. Applied to taxonomic rank categories the results imply that except for the ancient branching of monocots from dicots, only adaptive radiations of plants on the family and genus level are important for host utilization among phytophagous beetles. These findings enable improvements in estimating host specificity and species richness through correction for phylogenetic relatedness between hosts and consideration of the host-specific fauna associated with monocots.     

Consequences of defoliation on phenological interaction between Epirrita autumnata and its host plant, Mountain Birch

1. Defoliation-induced changes in the budbreak phenology of Mountain Birches ( Betula pubescens ssp. tortuosa (Ledeb.) Nyman), and their effects on herbivore performance, were investigated during an outbreak of the spring-feeding geometrid, Epirrita autumnata (Bkh.).
2. Total defoliation (followed by refoliation) by E. autumnata larvae was required to achieve significant delaying of budbreak of Mountain Birch one year after damage.
3. Delayed budbreak has potentially deleterious effects on herbivore performance. However, E. autumnata larvae show phenotypic responses that increase the synchrony between larvae and leaves: high larval density, a prerequisite for severe defoliation, delays egg hatch in the following year.
4. Late hatching larvae performed as well on birches with delayed budbreak due to the previous year's total defoliation as did early hatching larvae on birches with earlier budbreak.     

Fire effects on insect herbivores in an oak savanna: the role of light and nutrients

Abstract.  1. Environmental heterogeneity created by prescribed burning provided the context for testing whether the distribution of an oak specialist (the lace bug, Corythuca arcuata ) could be explained by stoichiometric mismatches between herbivore and host plant composition.
2. Field observations showed that lace bug density was seven-fold higher in frequently burned than in unburned units.
3. Lace bug density did not increase with leaf nutrient concentrations, but was instead associated with higher light levels, higher concentrations of leaf carbon (C), lignin and total phenolics, and lower levels of cellulose. In addition, lace bugs reared on high-light leaves had higher levels of survivorship than those fed on low-light leaves.
4. Sampling restricted to full-sun leaves was used to test whether fire-related changes in leaf nitrogen (N) and phosphorus (P) concentrations have a secondary influence on lace bug success. This sampling provided only limited evidence for nutrient limitation, as decreases in leaf N and P were associated with an increase in lace bug mass but a decrease in density.
5. It is concluded that burning probably promotes lace bug population growth by increasing canopy openness, light penetration, and the availability of C-based metabolites, and thus simple stoichoimetric mismatches between herbivores and host plants are not of primary importance in this system.     

Opposing survivorship and fecundity effects of host phenology on the gall-forming aphid Hormaphis hamamelidis

Abstract  1. This study examined the synchronisation and roles of host and insect phenology for fundatrices of the gall-forming aphid Hormaphis hamamelidis , which forms galls on the leaves of the understorey tree Hamamelis virginiana .
2. Host and insect phenologies varied widely among years and individual host trees.
3. Most winter eggs (≈ 85%) hatched before budbreak and galls could be initiated but in 2 years there was no inherent risk for earlier hatching fundatrices in terms of greater mortality. Egg hatch was not synchronised tightly with host trees but instead seemed to occur well in advance of budbreak as a conservative, bet-hedging strategy to ensure the opportunity to form galls.
4. Fundatrices that formed galls later had higher fecundity, contrary to typical patterns for insects that feed on the first flush of leaves in temperate forests, where later herbivores have lower fecundity. Also, unlike many other systems, trees that had later budbreak had greater densities of galls.     

Effects of tree species diversity and genotypic diversity on leafminers and parasitoids in a tropical forest plantation

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The Impact of Hydrological Restoration on Benthic Aquatic Invertebrate Communities in a New Zealand Wetland

Drainage is a major disturbance affecting wetlands, as drains lower water tables and convert lentic habitats to lotic ones. Consequently, invertebrate communities in drained wetlands are likely to differ from those in unimpacted wetlands. This study investigated the effect of hydrological restoration on invertebrate communities in small drains in a New Zealand fen. Invertebrates were collected over 4 summers from 10 drains within the wetland, one of which was blocked as part of a restoration program. The sampling protocol thus represented a Before‐After Control‐Impact experiment. Invertebrate community composition varied over the 4 years, but variability was greatest in the manipulated drain before and after it was blocked. Relative abundance of the amphipod Paraleptamphopus decreased after blockage, whereas those of the midges Chironomus zelandicus and Tanypodinae increased. Relative abundances of these taxa in control sites were unchanged. Hydraulic restoration thus had a demonstrable impact on the invertebrate communities. The invertebrate community of the blocked drain was compared to that of natural wetlands in undisturbed catchments. Similarity was very low prior to drain blockage, but increased following drain blockage. Invertebrate communities in the restored drain were more similar to those of low pH wetlands than high pH wetlands. Given the goal of restoring the communities to those similar to natural conditions, this was a beneficial result. These results, coupled with studies that showed a decline in the cover of alien pasture grasses around the blocked drain, suggest that drain blockage represents a cost‐effective way of restoring wetland plant and aquatic invertebrate communities, especially where connectivity allows for the natural recruitment of these organisms into restored areas.     

The effects of some soil properties (soil nitrogen forms and moisture) on community structure in Mediterranean plant communities in the Black Sea region

Mediterranean ecosystems comprise the second biodiversity hotspot area after tropical rain forests and will be most affected by global climate change. Therefore, it is important to understand community dynamics for effective conservation in this region. We investigated the relationships between soil moisture, nitrogen forms and community structuring in Quercus ilexL., Erica arborea L. and Sarcopoterium spinosum (L.) Spach communities, representing different successional stages, distributed as Mediterranean enclaves on the Sinop Peninsula (Turkey). The soil moisture, ammonium, nitrate and nitrite content were measured seasonally. Differences in these abiotic parameters within and between communities over seasons were tested. Previously collected biotic data were then used to analyze the relationship between soil parameters and community structure. Significant differences in soil parameters within and between seasons were found within and between communities. Our results show that there are different relationships between soil moisture, nitrogen forms and community structure in Mediterranean plant communities representing different successional stages. Differentiation in vegetation structure during succession cause changes especially in the water and nitrate content of the soil, and these changes in turn affect the continuity of community structure in Mediterranean plant communities.     

Effects of sclerophylly and host choice on gall densities and herbivory distribution in an Australian subtropical forest

We hypothesize that upper canopies contrast with the understorey vegetation in gall density and survival because of greater mortality in the latter. We expect that high sclerophylly rates in the upper canopy leaves are a main cause of such pattern, and more important than other environmental traits, for instance related to altitude. The study was conducted in Lamington National Park, Queensland, Australia. Four independent vertical cylindrical transects through the forest canopy and one equivalent, horizontal understorey transect (20 m long and 1 m in diameter) were sampled at different altitude (300, 700, 900 and 1100 m above sea level) during two seasons. Total and damaged leaves were counted, and galls were collected and opened to determine if they were alive or what may have been the cause of death. Sclerophylly was estimated as specific foliar mass. Out of 72 sampled plant species, 29 presented galls, of which the greatest densities were concentrated on seven host species. A significant increase in sclerophylly with increasing canopy stratum height was observed, but had no direct effect on gall distribution. Total and live galls were more abundant in the canopy than in the understorey for all altitudes but 300 m, where a specific infestation on saplings of the canopy tree Arytera divaricata occurred. We found a positive gall establishment and survivorship in the upper canopy, along with a decrease in chewing herbivory, which resulted in decreasing risks of gall death by herbivory. An overall high sclerophylly rate in both canopy and understorey and the total number of galled host species suggest that the plant community studied is prone to gall establishment and evolution. However, a few hosts species with extreme infestation, such as A. divaricata and Ficus watkinsiana, override the community‐wide effect of sclerophylly. Our results emphasize how scarce and patchy distributed galls are.     

植物氮形态利用策略及对外来植物入侵性的影响

氮是影响外来植物入侵性的重要因素之一, 但相关研究多关注土壤氮水平的效应, 较少考虑氮形态的作用。为从土壤氮形态利用的角度阐释外来植物的入侵机制, 本文在植物氮形态利用策略分析的基础上, 综述了外来植物氮形态利用的偏好性及其对入侵性的影响。植物的氮形态利用策略有偏好性和可塑性两种, 这可能与植物对土壤氮形态特性的长期适应有关; 植物不仅可以对土壤氮形态做出响应, 而且还能改造土壤氮形态, 并对改变后的土壤氮形态做出反馈响应。很多外来植物入侵硝态氮占优势的干扰生境, 偏好硝态氮的外来植物与本地植物竞争硝态氮; 而偏好铵态氮的外来植物通过抑制土壤硝化作用, 营造铵态氮环境, 促进自身生长, 同时抑制偏好硝态氮的本地植物生长。然而, 植物氮形态利用策略不是一成不变的, 而是受多种生物和非生物因素共同作用影响的复杂过程, 今后应加强多因素交互作用对外来入侵植物氮形态利用策略的影响及机制研究, 更好地揭示氮形态利用策略, 尤其是氮形态利用的可塑性与外来植物入侵性的关系。     

The effect of herbivore density on host plant mediated interactions between two insects

Recent work suggests that foliar- and root-feeding insects can interact in a novel plus-minus fashion. However, the effects of herbivores may be different at different densities. This paper describes two laboratory experiments investigating the effect of increasing insect density on plant performance and the host-plant mediated interaction between a leaf-mining fly and root-feeding chafer larvae. Above- and below-ground insect herbivory decreased plant performance. The effects of the root feeder were, for some parameters, different between the two densities of chafer larvae. Leaf mining significantly decreased the performance of the chafer larvae, while root herbivory was found to increase the pupal weight (related to fecundity) of the leaf miner. The effects of root herbivory suggest that the relationships between herbivore density and plant performance may be curved as the greater chafer density had no significant increased effect on the plants or foliar-feeding insects.     

虫瘿多样性及其与寄主植物和环境间关系

虫瘿是自然界极常见的生物现象,凝聚着昆虫与植物间显著、复杂而密切的协同关系。本文主要阐述了致瘿昆虫的主要类群及其在植物上的致瘿部位、虫瘿外部形态、虫瘿发育过程、虫瘿内部结构、虫瘿寄主植物多样性以及虫瘿空间分布规律等,探讨了致瘿昆虫和寄主植物间相互关系,以及影响虫瘿空间分布的环境因素等。最后对目前虫瘿生物学存在的问题及以后的研究方向进行了讨论,以期为有害虫瘿的控制和有益虫瘿的开发与利用,以及致瘿昆虫与寄主植物间协同演化关系、致瘿昆虫的致瘿机理等研究奠定一定的理论基础。     

Herbivory and plant competition effects on the population dynamics of Naufraga balearica,a threatened narrow endemic species of the Balearic Islands

Naufraga balearica is a caespitose member of the Umbelliferae and is endemic to Majorca. We conducted a 12-year study in which permanent plots were monitored (each plot was divided into 100 cells) to assess (1) the natural population dynamics, (2) goat herbivory (1998–2003) and (3) plant competition. Using photographs and a G-test of independence, we analysed the frequency of cells in which a species was present and, for each annual transition, the frequency of cells where the species was stable, decreased or increased for each species and treatment (plots open to herbivores vs. closed plots). The natural dynamics of this threatened species showed a clear decline at the study site. Dactylis glomerata and Erodium reichardii were the species that appeared to have the greatest effect on the presence of N. balearica, especially in the closed plots. The competition effect of neighbouring species was not homogeneous throughout the 11 annual transitions analysed, suggesting that other factors affect the dynamics of the threatened species. Some herbivory pressure is beneficial to N. balearica because the species fluctuates more in the opened plots and the behaviour of the goats helps to control competing species, permitting the establishment of N. balearica.     

The functions of foliar nyctinasty: a review and hypothesis

Foliar nyctinasty is a plant behaviour characterised by a pronounced daily oscillation in leaf orientation. During the day, the blades of nyctinastic plant leaves (or leaflets) assume a more or less horizontal position that optimises their ability to capture sunlight for photosynthesis. At night, the positions that the leaf blades assume, regardless of whether they arise by rising, falling or twisting, are essentially vertical. Among the ideas put forth to explain the raison d'être of foliar nyctinasty are that it: (i) improves the temperature relations of plants; (ii) helps remove surface water from foliage; (iii) prevents the disruption of photoperiodism by moonlight; and (iv) directly discourages insect herbivory. After discussing these previous hypotheses, a novel tritrophic hypothesis is introduced that proposes that foliar nyctinasty constitutes an indirect plant defence against nocturnal herbivores. It is suggested that the reduction in physical clutter that follows from nocturnal leaf closure may increase the foraging success of many types of animals that prey upon or parasitise herbivores. Predators and parasitoids generally use some combination of visual, auditory or olfactory cues to detect prey. In terrestrial environments, it is hypothesised that the vertical orientation of the blades of nyctinastic plants at night would be especially beneficial to flying nocturnal predators (e.g. bats and owls) and parasitoids whose modus operandi is death from above. The movements of prey beneath a plant with vertically oriented foliage would be visually more obvious to gleaning or swooping predators under nocturnal or crepuscular conditions. Such predators could also detect sounds made by prey better without baffling layers of foliage overhead to damp and disperse the signal. Moreover, any volatiles released by the prey would diffuse more directly to the awaiting olfactory apparatus of the predators or parasitoids. In addition to facilitating the demise of herbivores by carnivores and parasitoids, foliar nyctinasty, much like the enhanced illumination of the full moon, may mitigate feeding by nocturnal herbivores by altering their foraging behaviour. Foliar nyctinasty could also provide a competitive advantage by encouraging herbivores, seeking more cover, to forage on or around non‐nyctinastic species. As an added advantage, foliar nyctinasty, by decreasing the temperature between plants through its effects on re‐radiation, may slow certain types of ectothermic herbivores making them more vulnerable to predation. Foliar nyctinasty also may not solely be a behavioural adaptation against folivores; by discouraging foraging by granivores, the inclusive fitness of nyctinastic plants may be increased.     

Body size and host range co-determine the altitudinal distribution of Neotropical tephritid flies

Aim We addressed the following questions: (1) Does tephritid body size tend to increase in species found at higher elevations, as predicted by Bergmann's rule? (2) Do tephritids conform to Rapoport's rule, so that species found at higher elevations tend to have broader altitudinal ranges? (3) More generally, how do body size and host range jointly affect the patterns of altitudinal distribution among Neotropical tephritid flies? Location The Mantiqueira mountain range, south‐eastern Brazil, at sites ranging from c. 700 to 2500 m a.s.l. Methods At each site we collected flower heads of all Asteraceae species to rear out endophagous immatures (from January to June in 1998 and 1999). We used structural equation models (SEM) to evaluate jointly the relationships between body size, host range and altitudinal distribution (range and mid‐point). Results Neotropical tephritid body size showed a negative relationship with altitudinal distribution. SE modelling showed no significant direct effect of body size on altitudinal range; however, it had significant indirect negative effects through host range and altitudinal mid‐point. The SE model was a good predictor of observed correlations and accounted for 84% of the variation in tephritid altitudinal range. Main conclusions The altitudinal range of flower‐head‐feeding tephritids is related to host range and is indirectly affected by body size via host range and altitudinal mid‐point. As predicted by Rapoport's rule, tephritids that occur at higher elevations also present wider altitudinal ranges. Bergmann's rule does not apply to Neotropical tephritids along a tropical elevational gradient, but rather its converse was found. Body size may determine host range by imposing a restriction upon large individuals using small flower heads. Host species turnover along the altitudinal gradient may be the main factor explaining the strong relationship between host range and insect elevational distribution.     

Asymmetric effects of a leaf‐chewing herbivore on aphid population growth

1. Plant responses to herbivory are often specific to the feeding guild of the attacking herbivore. These phytochemical responses to herbivore damage can affect herbivore performance and activity. Comprehensive studies on the ecological consequences of multi‐herbivore plant interactions are key to understanding plant–herbivore community dynamics. 2. This study examined how feeding damage by co‐occurring herbivores from separate feeding guilds, Myzus persicae (Sulzer), a sucking herbivore, and Leptinotarsa decemlineata (Say), a chewing herbivore, alter plant chemistry and indirectly affect herbivore performance. Performance was measured when each insect fed on plants individually, sequentially, or simultaneously in laboratory and field experiments. Phytohormone and glycoalkaloid content were measured for each feeding sequence to evaluate plant responses to herbivory by each guild. Mid‐season and end‐of‐season tuber yield were evaluated in the field study. 3. Damage by L. decemlineata negatively impacted M. persicae performance in both laboratory and field settings. Damage by M. persicae did not affect L. decemlineata performance in laboratory assays. However, L. decemlineata performance was positively affected by M. persicae herbivory in the field, but this effect was temporary. Although phytohormones and plant defences varied across treatments, they provide little resolution on interaction outcomes. 4. These results confirm that the presence of multiple feeding guilds on a single plant can affect these chewing and sucking herbivores differentially, but given the variability in our phytochemical analyses compared with other studies, the mechanism remains unclear. The study's findings show that aphids are negatively affected by chewing herbivores across systems, while aphids temporarily affected beetles positively.     

The development and effects of nitrogen deficiency in field-grown kiwifruit (Actinidia deliciosa) vines

The development and effects of nitrogen (N) deficiency in kiwifruit (Actinidia deliciosa Hayward) vines planted at three densities (25.0, 12.5 and 8.33 m² vine^–1) were examined in a long term (1982 to 1989) field experiment in which N was applied at rates from 0 to 200 kg N ha^–1 year^–1. The rate of applied N significantly affected leaf N concentrations every year from 1985 onwards, and the average leaf N concentrations declined throughout the experiment. Fruit N concentrations varied significantly with the level of applied N as early as 1986. The average fruit N concentrations varied strongly between years, and were inversely proportional to the fruit number (per m²), indicating that, after fruit set, growth of individual fruit was relatively insensitive to the vine N status. Effects of N supply on fruit yields resulted mostly from changes in fruit number (per m²). For vines planted at the high density, fruit yields responded significantly to the level of applied N each season from 1986 onwards. In any year, maximum fruit yields for vines planted at the high density were associated with leaf N concentrations (20 weeks after bud burst) of at least 1.8 mmol g^–1. For vines planted at low density, significant yield responses to the level of applied N were not recorded until 1988, and maximum yields in that year were associated with leaf N concentrations of at least 1.4 mmol g^–1. The delayed expression of effects of N deficiency on fruit yields for vines planted at low density appeared to follow a shift in partitioning of resources in favour of fruit growth. This shift in partitioning did not appear to be sustainable, and by 1989 the fruit yield response to applied N continued to the highest N level tested. In that year, the leaf N concentration associated with maximum yield was 1.8 mmol g^–1, the same as that recorded throughout the experiment for the vines planted at high density. In the last two seasons of the experiment, leaf necrosis developed extensively on vines receiving less than the highest rate of N. This necrosis appeared to be premature senescence resulting from N deficiency. Leaf chloride (Cl) concentrations increased significantly with increasing severity of N deficiency, but were never more than those associated with Cl toxicity. While N supply significantly affected fruit firmness immediately post-harvest, there were no significant effects on fruit firmness after 12–20 weeks storage.