Top-down effects of insect herbivores during early succession: influence on biomass and plant dominance

We tested the hypothesis that phytophagous insects would have a strong top-down effect on early successional plant communities and would thus alter the course of succession. To test this hypothesis, we suppressed above-ground insects at regular intervals with a broad-spectrum insecticide through the first 3 years of old-field succession at three widely scattered locations in central New York State. Insect herbivory substantially reduced total plant biomass to a similar degree at all three sites by reducing the abundance of meadow goldenrod, Solidago altissima. As a result, Euthamia graminifolia dominated control plots whereas S. altissima dominated insecticide-treated plots by the third year of succession. S. altissima is the dominant old-field herbaceous species in this region but typically requires at least 5 years to become dominant. Past explanations for this delay have implicated colonization limitation whereas our data demonstrate that insect herbivory is a likely alternative explanation. A widespread, highly polyphagous insect, the xylem-tapping spittlebug, Philaenus spumarius, appeared to be the herbivore responsible for the reduction in standing crop biomass at all three sites. Insect herbivory typically caused little direct leaf tissue loss for the ten plant species we examined, including S. altissima. Consequently, the amount of leaf area removed was not a reliable indicator of the influence of insect herbivory on standing crop biomass or on early succession. Overall, we found a strong top-down effect of insect herbivores on biomass at several sites, so our results may be broadly applicable. These findings run counter to generalizations that top-down effects of herbivores, particularly insects, are weak in terrestrial systems. These generalizations may not apply to insects, such as spittlebugs, that can potentially mount an effective defense (i.e., spittle) against predators and subsequently reach relatively high abundance on common plant species. Our results suggest that insect herbivory may play an important but often overlooked role during early old-field succession. Received: 26 December 1998 / Accepted: 3 April 1999     

Relationships between the effects of insect herbivory and sheep grazing on seasonal changes in an early successional plant community

Summary The effects of spring grazing by sheep and of natural levels of insect herbivory were studied in 1985 on a limestone field abandoned from arable land for four years. A split-plot design was adopted in which paddocks, arranged in Latin squares, were either left ungrazed or heavily grazed by sheep for ten days in April. Within each paddock plots were either sprayed regularly with Malathion-60 or untreated.Natural levels of insect herbivory, compared to the reduced levels in insecticide-treated plots, had effects of similar magnitude to those from the short burst of spring grazing. Many attributes of the grazed/insecticide-treated sward were either increased or decreased by a factor of two within a season. Both types of herbivore caused changes in the direction of plant succession as well as in its rate. Effects on early successional species were large and similar when caused by either type of herbivore. Effects on later successional species were often smaller, but also showed differences in the action of the two herbivore types, as did effects on sward height, species richness and total cover. The effects of sheep and insect herbivory were not always additive or in the same direction.The results suggest that manipulations of both mammal and insect herbivores may be powerful tools for directing changes in plant community composition.     

Localization of CRF immunoreactivity in the central nervous system of three vertebrate and one insect species

Summary By use of a specific antiserum against synthetic ovine corticotropin-releasing factor (CRF) in the peroxidase-antiperoxidase (PAP) immunocytochemical procedure (Vandesande and Dierickx 1976), CRF-like antigenic determinants were demonstrated in the central nervous system of a human fetus, the Wistar rat, the frog Rana ridibunda, and the American cockroach Periplaneta americana. The immunoreactive CRF-producing cells occur mainly in the nucleus paraventricularis of the rat, while in Rana ridibunda these cells occur in the nucleus praeopticus. Immunoreactive CRF-containing fibres were also visualized. Very clear CRF-immunoreactive products were observed in the brain as well as the corpora cardiaca (CC) and corpora allata (CA) of the cockroach Periplaneta americana. ACTH-immunoreactivity was also demonstrated in the brain-CC-CA complex of this insect. Double immunohistochemical staining (Vandesande 1983) also revealed that both the CRFand ACTH-like substances occur in different neurosecretory neurons and nerve fibres. These results suggest that the antigenic determinants of CRF are very similar in vertebrates and insects bespeaking their very long evolutionary history.     

Phenology and climate change: a long-term study in a Mediterranean locality

It is well documented that plant and animal phenology is changing in response to recent climate warming in the Palaearctic. However, few long-term data sets are currently available in the Mediterranean basin. The present study reports long-term temporal trends of several phenophases of 45 plants, 4 insects and 6 migratory insectivorous birds. Dynamic factor analyses performed with plant phenophases showed that most of those events occurring at spring and summer had common trends toward the advancement, especially since mid-1970s. However, during these last decades, insect phenology showed a steeper advance than plant phenology, suggesting an increase of decoupling of some plant–insect interactions, such as those between pollinators and flowers or herbivorous insects and their plant resources. All trans-Saharan bird species showed highly significant temporal trends in all studied phenophases (some of them covering most of the last century). In two species, the duration of stay is increasing due to both earlier arrivals and later departures. On the other hand, two wintering species showed a significant advancement in their arrival dates, while an opposite pattern were found for departures of each one. Only one of these species increased significantly its wintering stay. Bird departures were not related to local climate in any species. Our results demonstrate a key role of local temperatures behind interannual variability of most plant and insects phenophases, with especial emphasis in those occurring in spring and summer. Therefore, the common signal towards the advancement recorded since mid-1970s resulted from the recent rise in temperatures.     

Species temporal variability: epigaeic ant assemblages and management for abundance and scarcity

Summary Ant assemblages in five closely-spaced point habitats were studied over a three-year period in a subtropical area of South Africa. A total of 20,746 individuals in 56 species were sampled with pitfall traps. Rapid species turnover occurred during short periods after longer intervals of relatively constant species composition. There was annual cycling of total numbers of individuals. The temporal variance for total number of individuals within each habitat increased significantly with mean population size by a logarithmic regression. This indicates that population fluctuations are dependent upon the mean, with a distinctive relationship. The total number of species was cyclical, with annual and approximately half-yearly cycles. Each habitat nevertheless carried a characteristic mean number of species. The temporal variance of species richness showed a highly significant positive logarithmic relationship with the mean number of species. Pheidole megacephala (F.) accounted for 45% of all individuals. Its overall population showed annual cycles. Ph. megacephala population trend was the main determinant for seasonal variation of the whole assemblage in most habitats. Percentage dominance was positively linearly correlated with the variance and with mean population level. Although Ph. megacephala population variance increased with the mean, proportionate variability did not, indicating that high populations are no more variable than low populations. Five variability measures were compared with each other on ant assemblages containing only the common species (1st quartile) and then including the increasingly rare species (1st and 2nd, 1st, 2nd and 3rd, and all quartiles). Williamson's Variability Measure, using logarithmic or inverse sine transformations, gave variable results depending whether or not the rarer species were included in the analysis. The Coefficient of Fluctuation also gave variable results depending on assemblage size. The Coefficient of Variation and Proportionate Variability Measure gave similar and consistent results, indicating that common species are not more variable than rare ones. The rarer species are stenotopic and tend always to be rare, whereas the common species are eurytopic, highly competitive and demographically volatile. Control measures for common ants, and conservation of rare ants, should take cognizance of these trends.     

The Chitinase A from the baculovirus AcMNPV enhances resistance to both fungi and herbivorous pests in tobacco

Biotechnology has allowed the development of novel strategies to obtain plants that are more resistant to pests, fungal pathogens and other agents of biotic stress. The obvious advantages of having genotypes with multiple beneficial traits have recently fostered the development of gene pyramiding strategies, but less attention has been given to the study of genes that can increase resistance to different types of harmful organisms. Here we report that a recombinant Chitinase A protein of the Autographa californica nuclear polyhedrosis virus (AcMNPV) has both antifungal and insecticide properties in vitro. Transgenic tobacco plants expressing an active ChiA protein showed reduced damages caused by fungal pathogens and lepidopteran larvae, while did not have an effect on aphid populations. To our knowledge, this is the first report on the characterisation and expression in plants of a single gene that increases resistance against herbivorous pests and fungal pathogens and not affecting non-target insects. The implications and the potential of the ChiA gene for plant molecular breeding and biotechnology are discussed.     

Impact of the fungus Balansia henningsiana on Panicum agrostoides: frequency of infection,plant growth and reproduction,and resistance to pests

Summary The impact of the systemic fungus Balansia henningsiana (Clavicipitaceae) on the grass Panicum agrostoides was examined in field and greenhouse studies comparing infected and uninfected plants. Approximately one-half of all plants in three populations located in southern Indiana were infected. In field samples and greenhouse studies infected plants were significantly heavier than uninfected plants and produced significantly more tillers. Infection tended to suppress flowering but occasional asymptomatic tillers on infected plants produced healthy inflorescences. Although infected plants produced fewer inflorescences than uninfected plants as a proportion of total tillers, absolute numbers of inflorescences were similar in the two groups. Because other grasses infected by different species of Balansia and related fungi often are more resistant to insect damage, pest damage was quantified in one population. No differences between infected and uninfected plants were detected in levels of herbivory but infected plants had significantly less damage by the common leaf spot fungus Alternaria triticina. The results suggest that there is no selective disadvantage for plants infected by B. henningsiana.     

Variation in leaf trichomes and nutrients of Wigandia urens (Hydrophyllaceae) and its implications for herbivory

Leaf trichome variation was studied in a population of Wigandia urens (Hydrophyllaceae) in relation to water availability, diversity of herbivorous insects, and grazing rates. Plants of W. urens have glandular and urticant (stinging hairs) trichomes, and it is possible to distinguish two types of leaves: smooth leaves with only glandular trichomes and bristly leaves with both types of trichomes. Density of urticant trichomes was negatively correlated with weekly mean precipitation. Fourteen species of insects in five different orders feed on leaves of W. urens throughout the year. Grazing rates varied according to type and age of leaves. Grazing rates were lower in smooth than bristly young leaves but the rates were similar for mature leaves. Secondary metabolites, screened using qualitative methods, showed similar classes of compounds in both types of leaves. Bristly leaves had significantly higher concentrations of nitrogen, phosphorous and water and these nutritional differences were well correlated with the differences in grazing rates between bristly and smooth leaves.     

Capacity for sustained terrestrial locomotion in an insect: Energetics,thermal dependence,and kinematics

Summary The capacity for sustained, terrestrial locomotion in the cockroach. Blaberus discoidalis, was determined in relation to running speed, metabolic cost, aerobic capacity, and ambient temperature (T _a=15, 23, and 34°C; acclimation temperature=24°C). Steady-state thoracic temperature (T _tss) increased linearly with speed at each T _a.The difference between T _tss and T _awas similar at each experimental temperature with a maximum increase of 7°C. Steady-state oxygen consumption (VO_2ss) increased linearly with speed at each T _aand had a low thermal dependence (Q₁₀=1.0-1.4). The minimum cost of locomotion (the slope of the VO_2ss versus speed function) was independent of T _a.Cockroaches attained a maximal oxygen consumption (VO_2max). increased with T _afrom 2.1 ml O₂·g^-1·h^-1 at 15°C to 4.9 ml O₂·g^-1·h^-1 at 23°C, but showed no further increase at 34°C, VO_2max increased 23-fold over resting VO₂ at 23°C, 10-fold at 34°C, and 15-fold at 15°C. Endurance correlated with the speed at which VO_2max was attained (MAS, maximal aerobic speed). Temperature affected the kinematics of locomotion. compared to cockroaches running at the same speed, but higher temperatures (23–34°C), low temperature (15°C) increased protraction time, reduced stride frequency, and reduced stability by increasing body pitching. The thermal independence of the minimum cost of locomotion (C_min), the low thermal dependence of VO_2ss (i.e., y-intercept of the VO_2ss versus speed function), and a typical Q₁₀ of 2.0 for VO_2max combined to increase MAS and endurance in B. discoidalis when T _awas increased from 15 to 23°C. Exerciserelated endothermy enabled running cockroaches to attain a greater VO_2max, metabolic scope, and endurance capacity at 23°C than would be possible if T _tss remained equal to T _a. The MAS of B. discoidalis was similar to that of other arthropods that use trachea, but was 2-fold greater than ectotherms, such as salamanders, frogs, and crabs of a comparable body mass.Abbreviations T _a ambient temperature - T _t thoracic temperature - T _tss steady state thoracic temperature during exercise - T _trest thoracic temperature during rest - VO₂ oxygen consumption - VO_2rest oxygen consumption during rest - VO_2ss steady-state oxygen consumption during exercise - VO_2max maximal oxygen consumption; MAS maximum aerobic speed - C _min minimum cost of locomotion - t _end endurance time     

Effects of small mammals and vertebrate predators on vegetation in the Chilean semiarid zone

 We monitored the cover and seed bank response of shrubs, perennial herbs, and ephemeral plants to experimental exclusion of both the principal rodent herbivore, Octodon degus, and its vertebrate predators from 1989 to 1994 in a semiarid Chilean mediterranean site. Although both richness and species composition of the plant community at the study site were largely determined by abiotic factors (mainly rainfall and soil nutrients), predator and herbivore exclusion had significant effects on the relative abundance of several plant species. Experimental exclusion of herbivores was associated with increased cover of some shrubs and a perennial grass, and decreased cover and seed densities of several ephemerals, especially those exotic or restricted to areas underneath shrubs. Herbivores apparently reduced shrubs through browsing and indirectly affected herb cover and seed densities by opening up areas under shrubs and/or modifying physical and chemical conditions of the soil. Plant responses to predator exclusion were less clear. Nevertheless, higher cover of some shrubs and ephemerals in the presence of predators suggests tritrophic effects through changes in small mammal densities and/or foraging behavior. Received: 22 April 1996 / Accepted: 14 August 1996     

Directional specificity and encoding of muscle forces and loads by stick insect tibial campaniform sensilla,including receptors with round cuticular caps

In many systems, loads are detected as the resistance to muscle contractions. We studied responses to loads and muscle forces in stick insect tibial campaniform sensilla, including a subgroup of receptors (Group 6B) with unusual round cuticular caps in oval-shaped collars. Loads were applied in different directions and muscle contractions were emulated by applying forces to the tibial flexor muscle tendon (apodeme). All sensilla 1) were maximally sensitive to loads applied in the plane of joint movement and 2) encoded muscle forces but did not discharge to unresisted movements. Identification of 6B sensilla by stimulation of cuticular caps demonstrated that receptor responses were correlated with their morphology. Sensilla with small cuticular collars produced small extracellular potentials, had low thresholds and strong tonic sensitivities that saturated at moderate levels. These receptors could effectively signal sustained loads. The largest spikes, derived from sensilla with large cuticular collars, had strong dynamic sensitivities and signaled a wide range of muscle forces and loads. Tibial sensilla are apparently tuned to produce no responses to inertial forces, as occur in the swing phase of walking. This conclusion is supported by tests in which animals 'stepped' on a compliant surface and sensory discharges only occurred in stance.     

Response of Salvinia molesta to insect damage: changes in nitrogen,phosphorus and potassium content

Summary When adults and larvae of the weevil Cyrtobagous salviniae destroyed buds and tunnelled through rhizomes of Salvinia molesta, the plant responded by producting new growth which contained higher concentrations of nitrogen than in undamaged plants or in the older parent tissue of the same plant. Damage to leaves by the moth, Samea multiplicalis did not induce the same response. C. salviniae fed on the new growth and the higher nitrogen intake would have increased its reproductive capacity and enhanced its action as a biological control agent.Damage by both insects resulted in potassium leaching from the plant but no change in concentrations of phosphorus. The results support earlier suggestions that damage by C. salviniae might improve the qualtity of the host plant for this herbivore.     

Thesis,Antithesis and Synthesis in the Development of Modern Biology

A link scheme in biology is described between the University of Edinburgh and local state primary schools. Undergraduates reading biology or medicine are assigned for a year to schools where they help with biology projects. Visits by the students take place about once a week during term-time, and a wide variety of topics is covered. The university also sets aside materials and equipment that can be used as teaching aids when students visit schools. Participation is voluntary for both students and schools. The administration of the scheme and the benefits to all parties concerned are outlined.     

Invertebrate pests of canola and their management in Australia: a review

Abstract  In Australia, canola is subject to attack by at least 30 species of invertebrate pests, although the composition of this pest complex can vary between regions. Mites (e.g. the redlegged earth mite Halotydeus destructor and the blue oat mites Penthaleus spp.), lucerne flea ( Sminthurus viridis ) and false wireworms (e.g. the grey false wireworm Isopteron punctatissimus and the bronzed field beetle Adelium brevicorne ) are the major pests threatening the seedling establishment, whereas aphids (the cabbage aphid Brevicoryne brassicae , the turnip aphid Lipaphis erysimi and the green peach aphid Myzus persicae ), the native budworm ( Helicoverpa punctigera ), the diamondback moth ( Plutella xylostella ) and the Rutherglen bug ( Nysius vinitor ) can cause irregular and unpredictable damage to the flowering and podding plants. Current tactics of pest management for canola rely largely on the use of synthetic pesticides, but this single-technology approach is likely to incur negative effects on natural enemies and the risk of pest resistance. Thus, the sustainable production of canola requires integrated pest management (IPM) strategies, in which cultural control, crop resistance and biological control are used as important components, with chemical inputs applied only when absolutely needed to restrict pests from reaching economically damaging densities. Such IPM strategies should be built around a fundamental understanding of pest ecology at both regional and local farm levels and the integration of renewable technologies. Therefore, future research efforts need to be focused on the canola-cropping system, with a particular emphasis on the impact of pest species, natural enemies of the pests, varietal resistance to pests and the spatial ecology of pest species.     

Controlling wildlife fungal disease spread: in vitro efficacy of disinfectants against Batrachochytrium dendrobatidis and Mucor amphibiorum

Chytridiomycosis in amphibians, and mucormycosis in the platypus Ornithorhynchus anatinus and amphibians, are serious fungal diseases affecting these aquatic taxa. In Tasmania, Australia, the fungi that cause these diseases overlap in range along with Phytophthora cinnamomi (Pc), an invasive fungal plant pathogen. To identify disinfectants that may be useful to reduce anthropogenic spread of these fungi to uninfected wilderness areas, for example by bush walkers and forestry or fire-fighting operations, we tested 3 disinfectants and a fire-fighting foam against Mucor amphibiorum (Ma) and tested 1 disinfectant and the foam against Batrachochytrium dendrobatidis (Bd). Combining the present study with previous work we found Bd was more susceptible to all 4 chemicals than Ma. Phytoclean, a disinfectant used at 2 to 10% for 30 s to control Pc, killed cultures of Bd at 0.075% and Ma at 5%, when also applied for 30 s. The disinfectant F10sc was not effective against Ma at standard exposures, but previous work shows Bd is killed at 0.03% with a 1 min exposure. Path-X is effective against Bd at 0.001% with a 30 s exposure and killed Ma at 1% with a 5 min exposure. Forexpan S, a foam added to water at 0.1 to 1% to control forest fires, killed Bd but not Ma when used at 1% for 2 min. Therefore, Phytoclean and Path-X have broader efficacy, although Path-X has not been trialled against Pc. Interestingly a positive mating strain of Ma (from a platypus) was more resistant to disinfectants than a negative strain (from a frog). Current protocols against Pc that involve high concentrations (10%) of Phytoclean are likely to reduce spread of pathogenic wildlife fungi, which is important for protecting biodiversity.     

Macroalgae, nutrients and phase shifts on coral reefs: scientific issues and management consequences for the Great Barrier Reef

 Degradation of coral reefs often involves a “phase shift” from abundant coral to abundant macroalgae. This paper critically reviews the roles of nutrient increases in such phase shifts. I conclude that nutrient overloads can contribute to reef degradation, but that they are unlikely to lead to phase shifts simply by enhancing algal growth rates and hence allowing overgrowth of corals, unless herbivory is unusually or artificially low. Concentrations of dissolved inorganic nutrients are poor indicators of reef status, and the concept of a simple threshold concentration that indicates eutrophication has little validity. I discuss the significance and consequences of these assessments for reef management, focusing on the Great Barrier Reef, and conclude with some specific recommendations, including protection of herbivorous fishes, minimisation of terrestrial runoff, and protection of coastal reefs. Accepted: 13 August 1999