The nutritional ecology of larvae of Alsophila pometaria and Anisota senatoria feeding on early- and late-season oak foliage

The larvae of Alsophila pometaria (Harr.), feeding on the young foliage of oak, has a higher relative growth rate (RGR) and relative nitrogen accumulation rate (RNAR) than the larvae of Anisota senatoria (J. E. Smith), feeding on the mature foliage of oak. Although the young oak foliage is more efficiently digested by A. pometaria (higher AD's), it is not more efficiently assimilated and used for growth (no difference in ECI's). Thus, the higher growth rate of A. pometaria is due entirely to a higher consumption rate (RCR and RNCR). Young foliage is significantly higher in nitrogen and water than mature foliage, but phenol and tannin levels are comparable in young and old foliage. A. pometaria consumes the foliage of different oak species at the same rate, independent of nitrogen content, while A. senatoria increases its consumption rate in response to decreased nitrogen levels. As a result, the growth rate of A. pometaria is directly related to leaf nitrogen content, while the growth rate of A. senatoria is independent of leaf nitrogen. The two species of insects have digestive systems that are very similar biochemically, and that are well-designed for effective protein digestion. Tannins and phenols do not influence the nutrional indices of either species. We suggest that the major benefit of spring feeding is the availability of succulent, high-nitrogen foliage, and not the avoidance of high-tannin foliage. The spring feeder appears to have a feeding strategy that favors rapid growth at the expense of efficiency, while the late summer feeder has a strategy that favors efficiency over rate.

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Résumé Alimentées sur feuillage jeune de chêne, les chenilles d'Alsophila pometaria avaient un taux relatif de croissance (RGR) et un taux relatif d'accumulation d'azote (RNAR) plus élevés que les chenilles d'Anisota senatoria alimentées sur feuillage mûr de chêne. Bien que le jeune feuillage soit plus efficacement digéré par A. pometaria (AD plus élevé), il n'est pas assimilé et utilisé pour la croissance avec de meilleurs rendements (les ECI ne sont pas différents). Ainsi le taux de croissance plus élevé d'A. pometaria est dû entièrement à un taux de consommation plus important (RCR et RNCR). Le feuillage jeune est significativement plus riche en azote et en eau que le feuillage mûr, mais les niveaux de phénol et de tanins sont les mêmes. A pometaria consomme les feuilles de différentes espèces de chênes au même taux, indépendamment de la teneur en azote, tandis que A. senatoria accroît sa consommation en réponse à une diminution de la teneur en azote. Il en résulte que le taux de croissance d'A. pometaria dépend directement de la teneur en azote des feuilles, tandis que celui d'A. senatoria en est indépendant. Les systèmes digestifs des deux insectes sont biochimiquement semblables et sont efficaces pour la digestion des protéines. Les tanins et les phénols n'influent pas sur les indices nutritionnels de ces deux espèces. Nous estimons que le principal intérêt de l'alimentation printanière est la disponibilité en feuillage succulent, riche en azote, et non l'absence de feuilles à haute teneur en tanin. L'alimentation printanière semble correspondre à une strategie alimentaire qui favorise la croissance aux dépens de l'efficacité tandis que l'alimentation en fin d'été est une stratégie qui favorise l'efficacité sur la rapidité.

     

Herbivoran impact on phytoplankton community structure

An enclosure experiment was conducted to assess the effects of a zooplankton elimination on the structure of a phytoplankton community. Phytoplankton biomass and production were higher in grazer-free enclosures, while the productivity per unit biovolume was lower. Exclusion of zooplankton favoured the majority of algal species, especially chrysophyceans (Dinobryon spp.) and the diatom Rhizosolenia, while mucilagineous green-algae were disfavoured. Middle sized algae (ESD 15–50 µm) and those with the largest Surface Area/Volume ratio were proportionally most favoured by the elimination of grazers.These differences in phytoplankton community structure are discussed in relation to effects of direct selective grazing and nutrient recycling by zooplankton. Some differences, as the immediate positive response of Dinobryon and Rhizosolenia, are probably caused by grazing release, while others, e.g. the response of mucilagineous species, might be caused by changed competitive relationships between the algae.     

Physiological and growth responses of Centaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability

Summary Centaurea maculosa seedlings were grown in pots to study the effects of root herbivory by Agapeta zoegana L. (Lep.: Cochylidae) and Cyphocleonus achates Fahr. (Col.: Curculionidae), grass competition and nitrogen shortage (each present or absent), using a full factorial design. The aims of the study were to analyse the impact of root herbivory on plant growth, resource allocation and physiological processes, and to test if these plant responses to herbivory were influenced by plant competition and nitrogen availability. The two root herbivores differed markedly in their impact on plant growth. While feeding by the moth A. zoegana in the root cortex had no effect on shoot and root mass, feeding by the weevil C. achates in the central vascular tissue greatly reduced shoot mass, but not root mass, leading to a reduced shoot/root ratio. The absence of significant effects of the two herbivores on root biomass, despite considerable consumption, indicates that compensatory root growth occurred. Competition with grass affected plant growth more than herbivory and nutrient status, resulting in reduced shoot and root growth, and number of leaves. Nitrogen shortage did not affect plant growth directly but greatly influenced the compensatory capacity of Centaurea maculosa to root herbivory. Under high nitrogen conditions, shoot biomass of plants infested by the weevil was reduced by 30% compared with uninfested plants. However, under poor nitrogen conditions a 63% reduction was observed compared with corresponding controls. Root herbivory was the most important stress factor affecting plant physiology. Besides a relative increase in biomass allocation to the roots, infested plants also showed a significant increase in nitrogen concentration in the roots and a concomitant reduction in leaf nitrogen concentration, reflecting a redirection of the nitrogen to the stronger sink. The level of fructans was greatly reduced in the roots after herbivore feeding. This is thought to be a consequence of their mobilisation to support compensatory root growth. A preliminary model linking the effects of these root herbivores to the physiological processes of C. maculosa is presented.     

Littorina mariae — a factor structuring low shore communities?

The role of littorinids in structuring communities is discussed. On hard substrates preferential grazing of the dominant algae often enables competitively inferior species to utilize the rock substrate. Investigations of these systems has led to the proposal of a number of factors that should be analyzed in order to assess the effect of herbivores on community structure. Some of these factors have been investigated for Littorina mariae Sacchi & Rastelli. L. mariae is a micro-epiphytic grazer browsing the surface covering of epiphytes off the alga Fucus serratus (L.). The relationship between the host alga, F. serratus, and L. mariae is far more intricate than that between winkles and hard substrates as the alga itself is a dynamic resource. L. mariae is spatially and temporally linked to F. serratus. L. mariae is found almost exclusively on the alga and is positively attracted by extracts of the alga. The life history of the winkle is closely synchronized with that of the alga (the winter decrease of L. mariae populations is associated with the seasonal die-back of the host alga). On analysis of the factors considered important to assess herbivory, and examples of the effects of other epiphytic herbivores on algal success, it is suggested that L. mariae could potentially play an important role in structuring the community of it's host fucoid and that this may influence larger scale community structure. Experimental manipulations are proposed to evaluate these hypotheses.     

Community structure and abundance of insects in response to early‐season aphid infestation in wild cabbage populations

1. Changes in the arthropod community structure can be attributed to differences in constitutively expressed plant traits or those that change depending on environmental conditions such as herbivory. Early‐season herbivory may have community‐wide effects on successive insect colonisation of host plants and the identity of the initially inducing insect may determine the direction and strength of the effects on the dynamics and composition of the associated insect community. 2. Previous studies have addressed the effect of early infestation with a chewing herbivore. In the present study, the effect of early infestation was investigated with a phloem‐feeding aphid [Brevicoryne brassicae L. (Hemiptera, Aphididae)] on the insect community associated with three wild cabbage (Brassica oleracea L.) populations, which are known to differ in defence chemistry, throughout the season in field experiments. 3. Aphid infestation had asymmetric effects on the associated insect community and only influenced the abundance of the natural enemies of aphids, but not that of chewing herbivores and their natural enemies. The effect size of aphid infestation further depended on the cabbage population. 4. Aphid feeding has been previously reported to promote host‐plant quality for chewing herbivores, which has been attributed to antagonism between the two major defence signalling pathways controlled by the hormones salicylic acid (SA) and jasmonic acid (JA), respectively. Our results show no effects of early infestation by aphids on chewing herbivores, suggesting the absence of long‐term JA–SA antagonism. 5. Investigating the effects of the identity of an early‐season coloniser and genotypic variation among plant populations on insect community dynamics are important in understanding insect–plant community ecology.     

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.     

Life‐history constraints in grassland plant species: a growth‐defence trade‐off is the norm

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade‐off possibilities. The competition‐defence hypothesis posits a trade‐off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth‐defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth‐defence trade‐off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life‐history groups and within the majority of individual sites. Thus, a growth‐defence trade‐off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.     

Optimizing canopy‐forming algae conservation and restoration with a new herbivorous fish deterrent device

The role of herbivorous fish in threatening marine forests of temperate seas has been generally overlooked. Only recently, the scientific community has highlighted that high fish herbivory can lead to regime shifts from canopy‐forming algae to less complex turf communities. Here, we present an innovative herbivorous fish deterrent device (DeFish), which can be used for conservation and restoration of marine forests. Compared to most traditional fish exclusion systems, such as cages, the DeFish system does not need regular cleaning and maintenance, making it more cost‐efficient. Resistance of DeFish was tested by installing prototypes at different depths in the French Riviera and in Montenegro: more than 60% of the devices endured several years without maintenance, even if most of them were slightly damaged in the exposed site in Montenegro. The efficacy of DeFish in limiting fish herbivory was tested by an exclusion experiment on Cystoseira amentacea in the French Riviera. In a few months, the number of fish bite marks on the seaweed was decreased, causing a consequent increase in algal length. The device here presented has been conceived for Mediterranean canopy‐forming algae, but the same concept can be applied to other species vulnerable to fish herbivory, such as kelps or seagrasses. In particular, the DeFish design could be improved using more robust and biodegradable materials. Innovative engineering systems, such as DeFish, are expected to become useful tools in the conservation and restoration of marine forests, to complement other practices including active reforestation, herbivore regulation, and regular monitoring of their status.