Changes in the Dynamics of Foliar N Metabolites in Oak Saplings by Drought and Air Warming Depend on Species and Soil Type

Climate change poses direct or indirect influences on physiological mechanisms in plants. In particular, long living plants like trees have to cope with the predicted climate changes (i.e. drought and air warming) during their life span. The present study aimed to quantify the consequences of simulated climate change for foliar N metabolites over a drought-rewetting-drought course. Saplings of three Central European oak species (i.e. Quercus robur, Q. petraea, Q. pubescens) were tested on two different soil types (i.e. acidic and calcareous). Consecutive drought periods increased foliar amino acid-N and soluble protein-N concentrations at the expense of structural N in all three oak species. In addition, transient effects on foliar metabolite dynamics were observed over the drought-rewetting-drought course. The lowest levels of foliar soluble protein-N, amino acid-N and potassium cation with a minor response to drought and air warming were found in the oak species originating from the driest/warmest habitat (Q. pubescens) compared to Q. robur and Q. petraea. Higher foliar osmolyte-N and potassium under drought and air warming were observed in all oak species when grown on calcareous versus acidic soil. These results indicate that species-specific differences in physiological mechanisms to compensate drought and elevated temperature are modified by soil acidity.     

Colonization of eucalypts in New Zealand by Australian insects

Since the 1860s, Australian insects have steadily colonized eucalypts in New Zealand. The rate of colonization has increased markedly over the last two decades. This increase may be related to increasing trade between the two countries. Currently there are 26 specialist eucalypt insect species and approximately 31 polyphagous insect species that can feed on Eucalyptus in New Zealand. The specialist eucalypt insects endemic to Australia have generally caused more damage than polyphagous or native insects. Eucalypt‐specific insects are dominated by sap sucking bugs, particularly psyllids, and defoliating Coleoptera and Lepidoptera. In some cases the major insect pest species have been those that are only occasional pests in Australia, for example Gonipterus scutellatus, Ctenarytaina eucalypti, Eriococcus coriaceus and Phylacteophaga froggatti. Some important insect pests have been rare, or even not described from Australia, prior to their appearance as a pest in New Zealand, for example Paropsis charybdis and Ophelimus eucalypti. Invading eucalypt insects are more likely to establish in the Auckland region than anywhere else in New Zealand.     

Strength and variability of postmating reproductive isolating barriers between four European white oak species

The identification and quantification of the relative importance of reproductive isolating barriers is of fundamental importance to understand species maintenance in the face of interspecific gene flow between hybridising species. Yet, such assessments require extensive experimental fertilisations that are particularly difficult when dealing with more than two hybridising and long-generation-time species such as oaks. Here, we quantify the relative contribution of four postmating reproductive isolating barriers consisting of two prezygotic barriers (gametic incompatibility, conspecific pollen precedence) and two postzygotic barriers (germination rate, early survival) from extensively controlled pollinations between four oak species (Quercus robur, Quercus petraea, Quercus pubescens and Quercus pyrenaica) that have been shown to frequently hybridise in natural populations. We found high variation in the strength of total reproductive isolation between species, ranging from total reproductive isolation to advantage toward hybrid formation. As previously found, Q. robur pollen was unable to fertilise Q. petraea due to a strong reproductive isolating mechanism. On the contrary, Q. pubescens pollen was more efficient at fertilising Q. petraea than conspecific pollen. Overall, prezygotic barriers contribute far more than postzygotic barriers to isolate species reproductively, suggesting a role for reinforcement in the development of prezygotic barriers. Conspecific pollen precedence reduced hybrid formation when pollen competition was allowed; however, presence of conspecific pollen did not totally prevent hybridization. Our results suggest that pollen competition depends on multiple ecological and environmental parameters, including species local abundance, and that it may be of uppermost importance to understand interspecific gene flow among natural multispecies populations.     

Distinct male reproductive strategies in two closely related oak species

Reproductive strategies of closely related species distributed along successional gradients should differ as a consequence of the trade‐off between competition and colonization abilities. We compared male reproductive strategies of Quercus robur and Q. petraea, two partly interfertile European oak species with different successional status. In the studied even‐aged stand, trees of the late‐successional species (Q. petraea) grew faster and suffered less from intertree competition than trees of the early‐successional species (Q. robur). A large‐scale paternity study and a spatially explicit individual‐based mating model were used to estimate parameters of pollen production and dispersal as well as sexual barriers between species. Male fecundity was found to be dependent both on a tree's circumference and on its environment, particularly so for Q. petraea. Pollen dispersal was greater and more isotropic in Q. robur than in Q. petraea. Premating barriers to hybridization were strong in both species, but more so in Q. petraea than in Q. robur. Hence, predictions based on the competition–colonization trade‐off are well supported, whereas the sexual barriers themselves seem to be shaped by colonization dynamics.     

Tree species identity and forest composition affect the number of oak processionary moth captured in pheromone traps and the intensity of larval defoliation

1. Pure forests are often seen as being more prone to damage by specialist pest insects than mixed forests, and particularly mixed forests associating host and nonhost species. We addressed the effect of tree diversity on oak colonization and defoliation by a major specialist pest, the oak processionary moth (OPM)
2. We quantified the number of male OPM moths captured and larval defoliation in pure stands of two oak host species (Quercus robur and Quercus petraea) and in mixed stands associating the two oak species or each oak species with another nonhost broadleaved species. We conducted two complementary studies to test the effect of host species and stand composition: (i) we used pheromone trapping to compare the number of males OPM captured throughout the distribution of oak hosts in France and (ii) we noted the presence of OPM nests and estimated defoliation in mature forests of north‐eastern France.
3. Oak species and stand composition significantly influenced the number of male OPM captured and defoliation by OPM larvae. Quercus petraea was consistently more attractive to and more defoliated by OPM than Q. robur. Both oak trees were attacked more in pure stands than in mixed stands, in particular mixed stands associating oaks with another (nonhost) broadleaved species.
4. The results of the present study support the view that mixed forests are more resistant to specialist pest insects than pure stands, and also indicate that this trend depends on forest composition. Our study provides new insights into OPM ecology and has potential implications for forest management, including the management of urban forests where OPM causes serious human health issues.

     

Differential effects of elevated CO2 on acorn density, weight, germination, and predation among three oak species in a scrub-oak forest

Much research on the effects of elevated CO₂ on forest trees has focused on quantitative changes in photosynthesis, secondary chemistry, and plant biomass. However, plant fitness responses to rising CO₂ should also include quantitative measures of reproduction, since most forest systems are recruitment limited. Until now, it has proved very difficult to grow forest trees to sexual maturity in a CO₂‐enriched environment. This paper is the first of its kind to address the effects of elevated CO₂ on the reproduction of hardwood trees in a natural forest. Beginning in 1996, scrub‐oak vegetation, predominantly three species of scrub‐oaks, Quercus myrtifolia, Q. chapmanii, and Q. geminata, were grown inside eight chambers with elevated CO₂ (704 parts per million (ppm)) and eight with ambient CO₂ (379 ppm) at Kennedy Space Center, Florida. In elevated CO₂, acorn production increased significantly for the dominant species Q. myrtifolia and for Q. chapmanii, but it did not increase for the subdominant, Q. geminata. Acorn weight, germination rate, and predation by weevils were unaffected by CO₂. Thus, recruitment of some forest tree species into the Florida scrub‐oak community is likely to be accelerated in an atmosphere of increased CO₂. However, because the acceleration of recruitment differs among species, over the long term, Q. myrtifolia and Q. chapmanii will be favored over Q. geminata and this is likely to change patterns of species diversity.     

Regeneration of oaks (Quercus robur/Q. petraea) and three other tree species during long-term succession after catastrophic disturbance (windthrow)

In broadleaved temperate forests in Europe, oak (Quercus robur/Q. petraea) regeneration is reported to be weak or absent. However, most work concern seedlings or saplings, studied relatively few years. We studied a Picea abies stand in Sweden, windthrown and logged (all stems harvested) in 1969, testing the hypothesis that oaks regenerate in the long term among competing tree species after catastrophic disturbance. In 2011, after 40 years/growth seasons, we recorded live and dead trees in the new stand and investigated the surroundings, competition, and succession. The following trees, up to 26 m tall, colonized: Sorbus aucuparia, Betula pendula/B. pubescens, Fagus sylvatica, Q. robur/Q. petraea, and Corylus avellana (a shrub). Betula dominated, and only Fagus was regenerating in 2011. Sorbus had produced most of the dead trees, mainly or partly through intraspecific competition. In the stand, compared to the surroundings, Quercus, Picea, and Alnus glutinosa were under-represented, and Sorbus, Betula, and Fagus were over-represented. Yet, the density of Quercus was far from negligible; 48 large trees/ha. Most of the oaks (74 %) were co-dominant trees and many grew near Sorbus. Thus, oaks can survive and grow fast among pioneer trees and browsing animals, a conclusion which is supported by the literature (nine studies identified). High mortality of Sorbus is part of a long-term succession, where Fagus might come to dominate. However, Quercus likely will persist, in low density. We propose three key traits contributing to long-term persistence of Q. robur/Q. petraea in European temperate forests: long life span, ecological plasticity, and resistance to disturbances.     

Ambrosiodmus rubricollis (Eichhoff) (Coleoptera; Curculionidae; Scolytinae) associated with young Tasmanian blue gum trees

Eucalypts are among the most widely planted forest trees in the world, and outside their native Australian range, the main arthropod pests are sap‐sucking insects, defoliators, gall‐making insects and xylophagous beetles. We report on a new association between a polyphagous wood‐boring beetle and Tasmanian blue gum (Eucalyptus globulus Labill.) in Central Portugal. Unidentified wood‐boring insects were found attacking two three‐year‐old E. globulus trees showing signs of decline among otherwise healthy trees in a commercial plantation, in June 2018. Declining trees presented dead twigs and branches, and recently developed epicormic sprouts evident on the trunks. Insects emerging from logs were identified as Ambrosiodmus rubricollis (Eichhoff), a species native to eastern and southern Asia, with the taxonomic identification validated by molecular analysis. To our knowledge, this is the first record of A. rubricollis associated with a eucalypt species worldwide. It is not clear whether the beetles played any significant role on the decline of the trees, but Ambrosiodmus may be potential pests for several tree and shrub species in Europe, as these beetles can transport pathogenic fungi.     

Friend or foe? The role of leaf-inhabiting fungal pathogens and endophytes in tree-insect interactions

Trees are large organisms that structure forest ecosystems by providing an environment for an enormous diversity of animal, microbial and plant species. As these species use trees as their common hosts, many are likely to interact with each other directly or indirectly. From studies on herbaceous plant species we know that microbes can affect the interaction of plants with herbivorous insects, for example via changes in plant metabolite profiles. The consequences of fungal colonization for tree-insect interactions are, however, barely known, despite the importance of these ecological communities. In this review we explore the interaction of leaf-inhabiting pathogenic and endophytic fungi with trees and the consequences for tree-living insect herbivores. We discuss molecular, physiological, chemical, biochemical and ecological aspects of tree-fungus interactions and summarize the current knowledge on the direct and indirect effects of tree-inhabiting fungi on insect herbivores.Our mechanistic understanding of the tripartite interaction of trees with leaf-inhabiting fungi and insect herbivores is still in its infancy. We are currently facing substantial drawbacks in experimental methodology that prevent us from revealing the effect of one single fungal species on a particular insect herbivore species and vice versa. Future studies applying a versatile toolbox of modern molecular, chemical analytical and ecological techniques in combined laboratory and field experiments will unequivocally lead to a better understanding of fungus-tree-insect interactions.     

Season and light affect constitutive defenses of understory shrub species against folivorous insects

Understory shrubs contribute to overall species diversity, providing habitat and forage for animals, influence soil chemistry and forest microclimate. However, very little is known about the chemical defense of various shrub species against folivorous insects. Using six shrub species, we tested how seasonal changes and light conditions affect their constitutive defense to insect damage. We monitored leaf perforation, concentrations of total phenols, condensed tannins, nitrogen (N), and total nonstructural carbohydrates (TNC). Leaf damage caused by insects was low in Sambucus nigra, Cornus sanguinea, and Frangula alnus, intermediate in Corylus avellana and Prunus serotina, and high in Prunus padus. Leaves of all the species, when growing in high light conditions, had high concentrations of defense metabolites. Except for C. avellana, leaves of the other shrub species growing in full sun were less injured than those in shade. This may be due to higher concentrations of defense metabolites and lower concentrations of nitrogen. Similar patterns of the effects of light on metabolites studied and N were observed for leaves with varying location within the crown of individual shrubs (from the top of the south direction to the bottom of the north), as for leaves from shrubs growing in full sun and shrubs in the shade of canopy trees. A probable cause of the greater damage of more sunlit leaves of C. avellana was the fact that they were herbivorized mostly by Altica brevicollis, a specialist insect that prefers plant tissues with a high TNC level and is not very sensitive to a high level of phenolic compounds.     

Effects of an invasive alien tree on the diversity and temporal dynamics of an insect assemblage on an oceanic island

Native vegetation is frequently replaced by alien plants on isolated oceanic islands. The effects of such replacements by invasive plants on the diversity and temporal dynamics of island-endemic insects remain unclear. We examined flying insect communities using Malaise traps on the small island of Nishi-jima in the oceanic Ogasawara Archipelago in the northwestern Pacific. On the island, an alien tree, Casuarina equisetifolia, has become dominant, occupying 57.3?% of the vegetation area. The species richness, composition, and abundance of pollinators (bees), predators (wasps), and wood-boring beetles (cerambycids, mordellids, and elaterids) were compared in each summer season of 4?years among three vegetation types: C. equisetifolia forest, natural forest, and grassland. In the traps, 82.3?% of species captured were endemic to the archipelago. The grassland harbored the highest species richness of native bees and wasps, whereas the natural forest had the highest species richness of native wood-boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups. Principal response curves indicated that differences in species composition among the three vegetation types were consistent through time for all insect groups. Most insect species were more abundant in natural forest or grassland than in C. equisetifolia forest. Standard deviations in both the numbers of individuals and species estimated under a Bayesian framework suggested that annual fluctuations of abundance and species density were similar among vegetation types (except for elaterid abundance). Therefore, replacement by C. equisetifolia has likely altered insect species composition but has not necessarily dramatically affected the temporal dynamics of insect assemblages on the island.     

Potential invasion of China by exotic insect pests associated with tree seeds

A total of 39 insect species, mostly seed chalcids in the genus Megastigmus (Hymenoptera), but also midges (Diptera), are listed as potential seed-borne invaders of Chinese conifers. Although the number of native seed insects per conifer genus does not differ between China and other biogeographical regions, there are significantly fewer seed insects associated with each conifer genus in China than potential invaders. The eventual success of the invaders is likely to depend on the presence of native Chinese conifers that are congeneric with the original host, or on the presence of the original host as an exotic. When a substantial entomofauna is already associated with cones, competition for seed resources may limit the potential impact of invaders because seed insects are usually the last organisms to colonize the cone. A survey of 26 fir species, both native and introduced to Europe, showed that overall seed infestation by five species of exotic chalcids is negatively correlated to levels of damage by native insects, except on the original hosts of the chalcids. Similar patterns are hypothesized for native firs, spruces, Douglas firs, and larches in China. Uncontrolled importation of seeds and nuts of broad-leaved trees could also facilitate the introduction of seed chalcids, seed bruchids, tortricid moths and nut weevils into China. Only six species of seed chalcids are present in China, out of the 72 known to attack broad-leaved seeds over the world.     

A molecular and genomic reference system for conifer defence against insects

Insect pests are part of natural forest ecosystems contributing to forest rejuvenation but can also cause ecological disturbance and economic losses that are expected to increase with climate change. The white pine or spruce weevil (Pissodes strobi) is a pest of conifer forests in North America. Weevil–host interactions with various spruce (Picea) species have been explored as a genomic and molecular reference system for conifer defence against insects. Interactions occur in two major phases of the insect life cycle. In the exophase, adult weevils are free‐moving and display behaviour of host selection for oviposition that is affected by host traits. In the endophase, insects live within the host where mobility and development from eggs to young adults are affected by a complex system of host defences. Genetic resistance exists in several spruce species and involves synergism of constitutive and induced chemical and physical defences that comprise the conifer defence syndrome. Here, we review conifer defences that disrupt the weevil life cycle and mechanisms by which trees resist weevil attack. We highlight molecular and genomic aspects and a possible role for the weevil microbiome. Knowledge of this conifer defence system is supporting forest health strategies and tree breeding for insect resistance.     

Spatial Pattern of Attacks of the Invasive Woodwasp Sirex noctilio,at Landscape and Stand Scales

Invasive insect pests are responsible for important damage to native and plantation forests, when population outbreaks occur. Understanding the spatial pattern of attacks by forest pest populations is essential to improve our understanding of insect population dynamics and for predicting attack risk by invasives or planning pest management strategies. The woodwasp Sirex noctilio is an invasive woodwasp that has become probably the most important pest of pine plantations in the Southern Hemisphere. Our aim was to study the spatial dynamics of S. noctilio populations in Southern Argentina. Specifically we describe: (1) the spatial patterns of S. noctilio outbreaks and their relation with environmental factors at a landscape scale; and (2) characterize the spatial pattern of attacked trees at the stand scale. We surveyed the spatial distribution of S. noctilio outbreaks in three pine plantation landscapes, and we assessed potential associations with topographic variables, habitat characteristics, and distance to other outbreaks. We also looked at the spatial distribution of attacked trees in 20 stands with different levels of infestation, and assessed the relationship of attacks with stand composition and management. We found that the spatial pattern of pine stands with S. noctilio outbreaks at the landscape scale is influenced mainly by the host species present, slope aspect, and distance to other outbreaks. At a stand scale, there is strong aggregation of attacked trees in stands with intermediate infestation levels, and the degree of attacks is influenced by host species and plantation management. We conclude that the pattern of S. noctilio damage at different spatial scales is influenced by a combination of both inherent population dynamics and the underlying patterns of environmental factors. Our results have important implications for the understanding and management of invasive insect outbreaks in forest systems.     

Effect of Tree Species and Mycorrhizal Colonization on the Archaeal Population of Boreal Forest Rhizospheres

Group 1.1c Crenarchaeota are the predominating archaeal group in acidic boreal forest soils. In this study, we show that the detection frequency of 1.1c crenarchaeotal 16S rRNA genes in the rhizospheres of the boreal forest trees increased following colonization by the ectomycorrhizal fungus Paxillus involutus. This effect was very clear in the fine roots of Pinus sylvestris, Picea abies, and Betula pendula, the most common forest trees in Finland. The nonmycorrhizal fine roots had a clearly different composition of archaeal 16S rRNA genes in comparison to the mycorrhizal fine roots. In the phylogenetic analysis, the 1.1c crenarchaeotal 16S rRNA gene sequences obtained from the fine roots formed a well-defined cluster separate from the mycorrhizal ones. Alnus glutinosa differed from the other trees by having high diversity and detection levels of Crenarchaeota both on fine roots and on mycorrhizas as well as by harboring a distinct archaeal flora. The similarity of the archaeal populations in rhizospheres of the different tree species was increased upon colonization by the ectomycorrhizal fungus. A minority of the sequences obtained from the mycorrhizas belonged to Euryarchaeota (order Halobacteriales).     

Herbivore and Fungal Pathogen Exclusion Affects the Seed Production of Four Common Grassland Species

Insect herbivores and fungal pathogens can independently affect plant fitness, and may have interactive effects. However, few studies have experimentally quantified the joint effects of insects and fungal pathogens on seed production in non-agricultural populations. We examined the factorial effects of insect herbivore exclusion (via insecticide) and fungal pathogen exclusion (via fungicide) on the population-level seed production of four common graminoid species (Andropogon gerardii, Schizachyrium scoparium, Poa pratensis, and Carex siccata) over two growing seasons in Minnesota, USA. We detected no interactive effects of herbivores and pathogens on seed production. However, the seed production of all four species was affected by either insecticide or fungicide in at least one year of the study. Insecticide consistently doubled the seed production of the historically most common species in the North American tallgrass prairie, A. gerardii (big bluestem). This is the first report of insect removal increasing seed production in this species. Insecticide increased A. gerardii number of seeds per seed head in one year, and mass per seed in both years, suggesting that consumption of flowers and seed embryos contributed to the effect on seed production. One of the primary insect species consuming A. gerardii flowers and seed embryos was likely the Cecidomyiid midge, Contarinia wattsi. Effects on all other plant species varied among years. Herbivores and pathogens likely reduce the dispersal and colonization ability of plants when they reduce seed output. Therefore, impacts on seed production of competitive dominant species may help to explain their relatively poor colonization abilities. Reduced seed output by dominant graminoids may thereby promote coexistence with subdominant species through competition-colonization tradeoffs.     

Pest management through tropical tree conservation

Tropical trees can provide various ecological services to adjacent agricultural environments, including maintaining and amplifying the numbers of beneficial insects. In Mexico, certain tree species harbor a diverse guild of hymenopteran parasitoids that attack pest fruit flies (Diptera: Tephritidae) and are at the same time sources of valuable hardwood timber. Indigenous trees and their associated fauna are slowly disappearing due to forest clearance and the expansion of crop monocultures. Here we explore the relationship among pest and non-pest fruit flies, their fruit-hosts and parasitoids in the context of mango orchards and surrounding patches of uncultivated vegetation and propose a novel mechanism to use these associations in favor of conservation purposes and pest management. Trees of conservation biological control interest are classified as: (1) parasitoid multiplier plants, species that serve as alternate hosts for key fruit fly pests when their commercial hosts are not available, but in which they are unusually vulnerable to parasitism; (2) parasitoid reservoir plants, native or introduced trees in whose fruits non-pest fruit flies serve as hosts to generalist parasitoids that are able to attack pest tephritids in other species of commercially grown fruit; and (3) pest-based parasitoid reservoir plants, native or introduced species that are not economically important locally, but which harbor fruit flies that would be pests in other circumstances and that serve as hosts for parasitoids of the important pests in the vicinity. Protection, multiplication and dissemination of such tree species has the potential to increase the number of naturally produced fruit fly parasitoids and could assist in the management of tephritid pests in areas where destruction of forests has impoverished the historical sources of fruit fly natural enemies. Tropical forest conservation may help resource-poor farmers reduce crop losses, increase biodiversity within fruit-growing regions and conserve native forests for both conservation purposes and commercial use of native hardwoods.     

Vector Affinity and Diversity of Geosmithia Fungi Living on Subcortical Insects Inhabiting Pinaceae Species in Central and Northeastern Europe

Fungi from the genus Geosmithia (Ascomycota: Hypocreales) are associated with bark beetles (Coleoptera: Scolytinae), though little is known about ecology, diversity, and distribution of these fungi across beetle and its host tree species. This study surveyed the diversity, distribution and vector affinity of Geosmithia isolated from subcortical insects that colonized trees from the family Pinaceae in Central and Northeastern Europe. Twelve Geosmithia species were isolated from 85 plant samples associated with 23 subcortical insect species (including 14 bark beetle species). Geosmithia community composition was similar across different localities and vector species; although the fungal communities associated with insects that colonized Pinus differed from that colonizing other tree species (Abies, Larix, and Picea). Ten Geosmithia species from four independent phylogenetic lineages were not reported previously from vectors feeding on other plant families and seem to be restricted to the vectors from Pinaceae only. We conclude that presence of such substrate specificity suggests a long and stable association between Geosmithia and bark beetles.     

The impact of arbuscular mycorrhizal fungi on tomato plant resistance against Tuta absoluta (Meyrick) in greenhouse conditions

The symbiotic relationship between plants and arbuscular mycorrhizal fungi (AMF) improves plant growth and increases its resistance to pests and diseases. Mycorrhizal fungi are among the specialized fungi associated with the rhizosphere and are completely dependent on plant organic carbon. In this research tomato, Solanum lycopersicum L. was used as the host plant to evaluate the interaction effects between inoculation of tomato plant with AMF and feeding of tomato leaf miner, Tuta absoluta (Meyrick). In addition, plant growth parameters and growth rate of insect were assessed. The mycorrhizal treatment included a mixture of four fungal species (Funneliformis mosseae, Rhizophagus intraradices, R. irregularis and Glomus iranicus). The results of the experiment showed that tomato plant roots were well colonized (66.29%) by AMF and there was a significant mutual relationship between the insects feeding on the plants and the fungi. Feeding by the insects on plants inoculated with the fungus increased percentage of colonization by AMF in plants infested with the insect as compared to the control plants. The results also indicated that growth parameters and phosphorus content of the plants inoculated with fungi significantly increased compared to the control group. Moreover, significantly lower growth rate and consumption index observed in the T. absoluta larvae were fed on the leaves of plants treated with AMF compared to leaves of plants not inoculated with AMF.