Regulation of forest defoliating insects through small mammal predation: reconsidering the mechanisms.

Population densities of forest defoliating insects may be regulated by small mammal predation on the pupae. When outbreaks do occur, they often coincide with warm, dry weather and at barren forest sites. A proposed reason for this is that weather and habitat affect small mammal population density (numerical response) and hence pupal predation. We propose an alternative explanation: weather and habitat affect small mammal feeding behaviour (functional response) and hence the outbreak risks of forest pest insects. We report results from laboratory and field-enclosure experiments estimating rates of pupal predation by bank voles (Myodes glareolus) on an outbreak insect, the European pine sawfly (Neodiprion sertifer), at different temperatures (15 and 20 °C), in different microhabitats (sheltered and non-sheltered), and with or without access to alternative food (sunflower seeds). We found that the probability of a single pupa being eaten at 20 °C was lower than at 15 °C (0.49 and 0.72, respectively). Pupal predation was higher in the sheltered microhabitat than in the open one, and the behaviour of the voles differed between microhabitats. More pupae were eaten in situ in the sheltered microhabitat whereas in the open area more pupae were removed and eaten elsewhere. Access to alternative food did not affect pupal predation. The results suggest that predation rates on pine sawfly pupae by voles are influenced by temperature- and habitat-induced variation in the physiology and behaviour of the predator, and not necessarily solely through effects on predator densities as previously proposed.

Geometrid outbreak waves travel across Europe.

We show that the population ecology of the 9- to 10-year cyclic, broadleaf-defoliating winter moth (Operophtera brumata) and other early-season geometrids cannot be fully understood on a local scale unless population behaviour is known on a European scale. Qualitative and quantitative data on O. brumata outbreaks were obtained from published sources and previously unpublished material provided by authors of this article. Data cover six decades from the 1950s to the first decade of twenty-first century and most European countries, giving new information fundamental for the understanding of the population ecology of O. brumata. Analyses on epicentral, regional and continental scales show that in each decade, a wave of O. brumata outbreaks travelled across Europe. On average, the waves moved unidirectionally ESE-WNW, that is, toward the Scandes and the Atlantic. When one wave reached the Atlantic coast after 9-10 years, the next one started in East Europe to travel the same c. 3000 km distance. The average wave speed and wavelength was 330 km year(-1) and 3135 km, respectively, the high speed being incongruous with sedentary geometrid populations. A mapping of the wave of the 1990s revealed that this wave travelled in a straight E-W direction. It therefore passed the Scandes diagonally first in the north on its way westward. Within the frame of the Scandes, this caused the illusion that the wave moved N-S. In analogy, outbreaks described previously as moving S-N or occurring contemporaneously along the Scandes were probably the result of continental-scale waves meeting the Scandes obliquely from the south or in parallel. In the steppe zone of eastern-most and south-east Europe, outbreaks of the winter moth did not participate in the waves. Here, broadleaved stands are small and widely separated. This makes the zone hostile to short-distance dispersal between O. brumata subpopulations and prevents synchronization within meta-populations. We hypothesize that hostile boundary models, involving reciprocal host-herbivore-enemy reactions at the transition between the steppe and the broadleaved forest zones, offer the best explanation to the origin of outbreak waves. These results have theoretical and practical implications and indicate that multidisciplinary, continentally coordinated studies are essential for an understanding of the spatio-temporal behaviour of cyclic animal populations.

Synergistic effects of methyl jasmonate treatment and propagation method on Norway spruce resistance against a bark-feeding insect.

Utilizing plants with enhanced resistance traits is gaining interest in plant protection. Two strategies are especially promising for increasing resistance against a forest insect pest, the pine weevil (Hylobius abietis): exogenous application of the plant defense hormone methyl jasmonate (MeJA), and production of plants through the clonal propagation method somatic embryogenesis (SE). Here, we quantified and compared the separate and combined effects of SE and MeJA on Norway spruce resistance to pine weevil damage. Plants produced via SE (emblings) and nursery seedlings (containerized and bare-root), were treated (or not) with MeJA and exposed to pine weevils in the field (followed for 3 years) and in the lab (with a non-choice experiment). Firstly, we found that SE and MeJA independently decreased pine weevil damage to Norway spruce plants in the field by 32-33% and 53-59%, respectively, compared to untreated containerized and bare-root seedlings. Secondly, SE and MeJA together reduced damage to an even greater extent, with treated emblings receiving 86-87% less damage when compared to either untreated containerized or bare-root seedlings in the field, and by 48% in the lab. Moreover, MeJA-treated emblings experienced 98% lower mortality than untreated containerized seedlings, and this high level of survival was similar to that experienced by treated bare-root seedlings. These positive effects on survival remained for MeJA-treated emblings across the 3-year experimental period. We conclude that SE and MeJA have the potential to work synergistically to improve plants' ability to resist damage, and can thus confer a strong plant protection advantage. The mechanisms underlying these responses merit further examination.

Insecticides suppress natural enemies and increase pest damage in cabbage.

Intensive use of pesticides is common and increasing despite a growing and historically well documented awareness of the costs and hazards. The benefits from pesticides of increased yields from sufficient pest control may be outweighed by developed resistance in pests and killing of beneficial natural enemies. Other negative effects are human health problems and lower prices because of consumers' desire to buy organic products. Few studies have examined these trade-offs in the field. Here, we demonstrate that Nicaraguan cabbage (Brassica spp.) farmers may suffer economically by using insecticides as they get more damage by the main pest diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), at the same time as they spend economic resources on insecticides. Replicated similarly sized cabbage fields cultivated in a standardized manner were either treated with insecticides according common practice or not treated with insecticides over two seasons. Fields treated with insecticides suffered, compared with nontreated fields, equal or, at least in some periods of the seasons, higher diamondback moth pest attacks. These fields also had increased leaf damage on the harvested cabbage heads. Weight and size of the heads were not affected. The farmers received the same price on the local market irrespective of insecticide use. Rates of parasitized diamondback moth were consistently lower in the treated fields. Negative effects of using insecticides against diamondback moth were found for the density of parasitoids and generalist predatory wasps, and tended to affect spiders negatively. The observed increased leaf damages in insecticide-treated fields may be a combined consequence of insecticide resistance in the pest, and of lower predation and parasitization rates from naturally occurring predators that are suppressed by the insecticide applications. The results indicate biological control as a viable and economic alternative pest management strategy, something that may be particularly relevant for the production of cash crops in tropical countries where insecticide use is heavy and possibly increasing.

Comparing Exogenous Methods to Induce Plant-Resistance Against a Bark-Feeding Insect.

Exogenous application of the plant hormone methyl jasmonate (MeJA) can trigger induced plant defenses against herbivores, and has been shown to provide protection against insect herbivory in conifer seedlings. Other methods, such as mechanical damage to seedlings, can also induce plant defenses, yet few have been compared to MeJA and most studies lack subsequent herbivory feeding tests. We conducted two lab experiments to: (1) compare the efficacy of MeJA to mechanical damage treatments that could also induce seedling resistance, (2) examine if subsequent insect damage differs depending on the time since induction treatments occurred, and (3) assess if these induction methods affect plant growth. We compared Scots pine (Pinus sylvestris) seedlings sprayed with MeJA (10 or 15 mM) to seedlings subjected to four different mechanical bark damage treatments (two different bark wound sizes, needle-piercing damage, root damage) and previous pine weevil (Hylobius abietis) damage as a reference treatment. The seedlings were exposed to pine weevils 12 or 32 days after treatments (early and late exposure, hereafter), and resistance was measured as the amount of damage received by plants. At early exposure, seedlings treated with needle-piercing damage received significantly more subsequent pine weevil feeding damage than those treated with MeJA. Seedlings treated with MeJA and needle-piercing damage received 84% less and 250% more pine weevil feeding, respectively, relative to control seedlings. The other treatments did not differ statistically from control or MeJA in terms of subsequent pine weevil damage. For the late exposure group, plants in all induction treatments tended to receive less pine weevil feeding (yet this was not statistically significant) compared to control seedlings. On the other hand, MeJA significantly slowed down seedling growth relative to control and all other induction treatments. Overall, the mechanical damage treatments appeared to have no or variable effects on seedling resistance. One of the treatments, needle-piercing damage, actually increased pine weevil feeding at early exposure. These results therefore suggest that mechanical damage shows little potential as a plant protection measure to reduce feeding by a bark-chewing insect.

Associating Land Cover Changes with Patterns of Incidences of Climate-Sensitive Infections: An Example on Tick-Borne Diseases in the Nordic Area.

Some of the climate-sensitive infections (CSIs) affecting humans are zoonotic vector-borne diseases, such as Lyme borreliosis (BOR) and tick-borne encephalitis (TBE), mostly linked to various species of ticks as vectors. Due to climate change, the geographical distribution of tick species, their hosts, and the prevalence of pathogens are likely to change. A recent increase in human incidences of these CSIs in the Nordic regions might indicate an expansion of the range of ticks and hosts, with vegetation changes acting as potential predictors linked to habitat suitability. In this paper, we study districts in Fennoscandia and Russia where incidences of BOR and TBE have steadily increased over the 1995-2015 period (defined as 'Well Increasing districts'). This selection is taken as a proxy for increasing the prevalence of tick-borne pathogens due to increased habitat suitability for ticks and hosts, thus simplifying the multiple factors that explain incidence variations. This approach allows vegetation types and strengths of correlation specific to the WI districts to be differentiated and compared with associations found over all districts. Land cover types and their changes found to be associated with increasing human disease incidence are described, indicating zones with potential future higher risk of these diseases. Combining vegetation cover and climate variables in regression models shows the interplay of biotic and abiotic factors linked to CSI incidences and identifies some differences between BOR and TBE. Regression model projections up until 2070 under different climate scenarios depict possible CSI progressions within the studied area and are consistent with the observed changes over the past 20 years.

Identifying climate-sensitive infectious diseases in animals and humans in Northern regions.

BACKGROUND: General knowledge on climate change effects and adaptation strategies has increased significantly in recent years. However, there is still a substantial information gap regarding the influence of climate change on infectious diseases and how these diseases should be identified. From a One Health perspective, zoonotic infections are of particular concern. The climate in Northern regions is changing faster than the global average. This study sought to identify climate-sensitive infectious diseases (CSIs) of relevance for humans and/or animals living in Northern regions. Inclusion criteria for CSIs were constructed using expert assessments. Based on these principles, 37 potential CSIs relevant for Northern regions were identified. A systematic literature search was performed in three databases using an explicit stepwise approach to determine whether the literature supports selection of these 37 potential CSIs. RESULTS: In total, 1275 nominated abstracts were read and categorised using predefined criteria. Results showed that arthropod vector-borne diseases in particular are recognised as having potential to expand their distribution towards Northern latitudes and that tick-borne encephalitis and borreliosis, midge-borne bluetongue and the parasitic infection fasciolosis can be classified as climate-sensitive. Many of the other potential CSIs considered are affected by extreme weather events, but could not be clearly classified as climate-sensitive. An additional literature search comparing awareness of climate influences on potential CSIs between 1997-2006 and 2007-2016 showed an increase in the number of papers mentioning effects of climate change. CONCLUSIONS: The four CSIs identified in this study could be targeted in a systematic surveillance programme in Northern regions. It is evident that climate change can affect the epidemiology and geographical range of many infectious diseases, but there were difficulties in identifying additional CSIs, most likely because other factors may be of equal or greater importance. However, climate-ecological dynamics are constantly under change, and therefore diseases may fall in or out of the climate-sensitive definition over time. There is increasing awareness in the literature of the effects of climate change on infectious diseases over time.

Methods to identify the prey of invertebrate predators in terrestrial field studies.

Predation is an interaction during which an organism kills and feeds on another organism. Past and current interest in studying predation in terrestrial habitats has yielded a number of methods to assess invertebrate predation events in terrestrial ecosystems. We provide a decision tree to select appropriate methods for individual studies. For each method, we then present a short introduction, key examples for applications, advantages and disadvantages, and an outlook to future refinements. Video and, to a lesser extent, live observations are recommended in studies that address behavioral aspects of predator-prey interactions or focus on per capita predation rates. Cage studies are only appropriate for small predator species, but often suffer from a bias via cage effects. The use of prey baits or analyses of prey remains are cheaper than other methods and have the potential to provide per capita predation estimates. These advantages often come at the cost of low taxonomic specificity. Molecular methods provide reliable estimates at a fine level of taxonomic resolution and are free of observer bias for predator species of any size. However, the current PCR-based methods lack the ability to estimate predation rates for individual predators and are more expensive than other methods. Molecular and stable isotope analyses are best suited to address systems that include a range of predator and prey species. Our review of methods strongly suggests that while in many cases individual methods are sufficient to study specific questions, combinations of methods hold a high potential to provide more holistic insights into predation events. This review presents an overview of methods to researchers that are new to the field or to particular aspects of predation ecology and provides recommendations toward the subset of suitable methods to identify the prey of invertebrate predators in terrestrial field research.

Causes behind insect folivory patterns in latitudinal gradients.

1.Adams and Zhang recently published one of the best studies so far of patterns of insect folivory along a latitudinal (climatic) gradient. They show clear negative trends in foliage loss in relation to temperature for certain groups of insect herbivores.2.Although their suggestion that the plant-herbivore interaction may be more important in cooler climates could be valid, they did not bring up the complementary explanation that interactions between predators and herbivores could also vary with climate. There are indications that insect natural enemies may respond more positively than insect herbivores to an increase in temperature. We argue that higher predator pressure in warmer climates may partly explain the patterns observed by Adams and Zhang.3.Synthesis.To further develop the important research concerning herbivory in a changing climate, both theoretically and empirically, plant ecologists and entomologists would mutually benefit from joining forces.