Disease ontogeny overshadows effects of climate and species interactions on population dynamics in a nonnative forest disease complex

Biotic threats to trees often arise from interactions among two or more species, frequently insects and fungi, that function together to defeat host defenses, secure resources and colonize new hosts. Feedbacks among plant enemies can have large effects on host population and disease dynamics, either by promoting stabilizing negative feedbacks or contributing to positive feedbacks that can destabilize populations and permit outbreaks. Feedbacks can be rapid and direct (e.g. within trees or among years) or can arise from slowly developing changes in host resource quantity or quality at the scale of forest stands or landscapes. Climate may also influence system dynamics by altering feedbacks within or among species or through density independent effects. We evaluated major drivers of population dynamics of beech bark disease (BBD), an important forest disease in eastern deciduous forests of North America, using data from 28 study sites in the eastern United States monitored for up to 14 yr between 1979 and 1992. Both primary causal agents of BBD – the introduced felted beech scale Cryptococcus fagisuga and native fungi Neonectria spp. – showed strong simple density dependence in all study populations. Surprisingly, densities of scale insects and fungi had little or no effects on population growth rates of the other, despite their habit of living in close physical relationships. For both insects and fungi, ecologically important features of the density dependent functions (slope, carrying capacity and density independent variance) were variable across sites. Climatic effects on density‐dependent functions (and scatter around them) were evident but generally weak and variable. The most striking predictor of patterns in density dependence was duration since establishment of BBD in the region. Apparently BBD alters forests over decades in ways that strengthen self‐regulation among causal agents without eliminating or even dramatically reducing host populations.     

Long-term Changes in Forest Carbon and Nitrogen Cycling Caused by an Introduced Pest/Pathogen Complex

Invasion of exotic forest pests and pathogens is a serious environmental problem for many forests throughout the world, and has been especially damaging to forests of eastern North America. We studied the impacts of an exotic pest/pathogen complex, the beech bark disease (BBD), in the Catskill Mountains of New York State, USA. In this region, BBD has caused a decline in the basal area of American beech (Fagus grandifolia Ehrh.) over the last 60 years and this decline has been accompanied by an increase in the basal area of sugar maple (Acer saccharum Marsh.). We studied the impacts of the BBD on carbon (C) and nitrogen (N) cycling using a series of stands that represented a sequence of disease impact and beech replacement by sugar maple. Our study showed that these long-term changes in tree species composition can lead to important changes in C and N cycling in the ecosystem, including an increase in litter decomposition, a decrease in soil C:N ratio, and an increase in extractable nitrate in the soil and nitrate in soil solution. Rates of potential net N mineralization and nitrification did not change across the BBD sequence, but the fraction of mineralized N that was nitrified increased significantly. Many of the observed changes in ecosystem function are larger in magnitude than those attributed to climate change or air pollution, suggesting that the impacts of invasive pests and pathogens on tree species composition could be one of the most important factors driving changes in C and N cycling in these forests in the coming decades.     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

Beech forest management does not affect the infestation rate of the beech scale Cryptococcus fagisuga across three regions in Germany

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Early-stage of invasion by beech bark disease does not necessarily trigger American beech root sucker establishment in hardwood stands

Two concomitant phenomena currently affect the dynamics of sugar maple-American beech (AB) stands in northeastern North America: beech bark disease (BBD), and increased AB understory density. Many studies suggest a causal link between the two phenomena, i.e., BBD favouring beech regeneration. But this link has yet to be experimentally demonstrated. To address the question, we compared regeneration composition between recently BBD-affected and -unaffected stands. A total of 109 stands were sampled; half were affected by BBD. Seedling and sapling density were assessed, together with the origin (seedling or sprout). While BBD affects stands in the eastern part of the study region, AB was observed in the understory across the entire region. No clear difference in AB sprout density between BBD-affected and -unaffected stands was observed while AB seedling density—as well as pooled AB seedling and sprout density were higher in unaffected stands. Findings suggests that BBD, in its early stage, is not a necessary trigger of AB understory establishment. Yet, AB sapling basal area generally was higher in stands affected by BBD, likely indicating a greater rate of AB understory development due to increased light availability beneath a more open crown canopy. That development can lead to AB understory dominance. This distinction—BBD not necessarily triggering AB root sucker establishment but favoring AB advance regeneration development—also questions the generalized perception that dense AB thickets necessarily originate from root suckers.     

Neighbourhood analyses of tree seed predation by introduced rodents in a New Zealand temperate rainforest

House mice Mus musculus and other introduced rodents represent a novel source of predation on tree seeds in New Zealand forests. In the northern temperate forests where these rodents are native, spatial and temporal variation in tree seed production can result in dramatic fluctuations in the distribution and abundance of seed predators, with subsequent feedbacks on the distribution and abundance of seedlings. We use neighbourhood models to examine variation in rodent predation on seeds of 4 tree species of the temperate rainforests of New Zealand as a function of 1) spatial variation in local canopy composition and 2) spatial and temporal variation in mouse activity. We placed seeds throughout mapped stands of mixed forests in alluvial valley bottoms and on elevated marine terraces in the Waitutu Forest, South Island. The risk of predation on seeds of 2 dominant canopy trees – rimu Dacrydium cupressinum and mountain beech Nothofagus solandri var . cliffortioides – peaked in neighbourhoods dominated by those species and by silver beech N. menziesii , particularly in a year of plentiful seed rain from these species. The risk of predation on rimu and beech seed was also related to measures of local mouse activity. These relationships suggest that the highest local abundance of mice was concentrated in rimu and beech neighbourhoods because of the food provided by seed rain from those trees. Predation on seed of miro Prumnopitys ferruginea , which is eaten by rats but not mice, was low in rimu neighbourhoods and where mouse activity was high. These patterns may reflect spatial segregation in the activity of rats versus mice within stands. Our results suggest that the spatial distribution of canopy trees translates into predictable patterns of variation in mouse activity and seed predation. Heterogeneity in rodent activity and seed predation within stands may have important implications for tree population dynamics.     

Magellanic woodpecker ( Campephilus magellanicus) abundance and foraging in Tierra del Fuego, Chile

The Magellanic woodpecker (Campephilus magellanicus) is a vulnerable and poorly studied bird in the sub-antarctic deciduous and evergreen beech (Nothofagus) forests of South America. On Tierra del Fuego island (Chile), we compared Magellanic woodpecker abundance and its foraging habitat in two forest types: pure N. pumilio and mixed forests composed by N. pumilio and N. betuloides, including managed and non managed stands. At a regional scale, abundance of woodpeckers was greater in landscapes including both forest types than in pure N. pumilio landscapes. When both forest types occurred together, woodpecker abundance did not differ between them. The number of trees with foraging signs was correlated with Magellanic woodpecker abundance and was also associated with N. betuloides and snag densities, but was not affected by forest management. Occurrence of pecking on foraging trees was greater in mixed Nothofagus than pure N. pumilio stands. Woodpeckers foraged disproportionately more on larger diameter and more decayed trees. Moreover, trees used for foraging were positively correlated with canopy cover and snag density and were negatively correlated with distance to nearby peatlands and beaver ponds. Direct observation revealed that the flying distance between trees was negatively correlated with proportion of trees with foraging signs. Woodpeckers chose trees that were visited before, suggesting a pattern of tree recognition within foraging territories.Communicated by F. Bairlein     

Macromoth community structure along a 95-year post-harvest chronosequence in managed forests of northwest Washington State (U.S.A.), with comparison to old growth forest communities

1. The effects of timber harvest in the moist coniferous forests of western North America are not well documented for ecologically important arthropods such as moths.
2. We assessed the response of macromoth community structure (abundance, sample size-corrected estimates of species richness and diversity, and overall community composition) to time since deforestation at 20 previously logged sites (1–95 years post-harvest), and compared the macromoth communities at these stands to four old growth stands.
3. As stand age increased following timber harvest, the number of macromoths captured in ultraviolet light traps increased and the relative abundance of dietary generalists declined, but sample size-corrected estimates of species richness and diversity did not vary. Macromoth community composition of the youngest stands (<10 years post-harvest) differed markedly from each other but converged soon thereafter.
4. Macromoth communities at old growth sites featured higher capture rates, lower dominance by dietary generalists, and higher sample size-corrected estimates of species richness and diversity than at previously logged sites. Community composition profiles for old growth sites differed from all previously logged sites, but the differences were subtle except in comparison to the youngest logged sites. None of the 188 species we sampled were old growth specialists.
5. Our results reveal dramatic initial impacts of deforestation on macromoth communities in moist coniferous forests of western North America. Such effects are largely reversed within two decades post-harvest but some effects persist for at least 95-years following logging.

     

Dieback of riparian alder caused by the Phytophthora alni complex: projected consequences for stream ecosystems

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Short‐term effect of selection cutting in boreal balsam fir forest on cerambycid and scolytid beetles

The ecology and management of boreal forest ecosystems are drawing greater attention worldwide as their importance is being increasingly recognized for carbon sequestration or for harbouring the world's largest remaining intact forests. Selection cuts have been introduced as a more socially acceptable silvicultural method to improve the maintenance of habitat structure and functions as they mimic aspects of boreal forest succession dynamics. Many studies have shown that selection cutting helps maintaining arthropod communities in mature forests, but few have examined the increased risks of damage by bark‐ and wood‐boring insects in boreal forests of eastern North America. We used multidirectional flight‐interception traps to quantify the response of these beetles to 25 and 40% selection cutting in a balsam fir–white birch forest of Québec, Canada. The abundance and species number of both cerambycid and scolytid beetles were 5–6 times larger in selectively cut stands than in controls the year following treatments. Analyses revealed that bark‐ and wood‐boring beetles’ response was mostly associated with increased canopy openness in selectively cut stands (and sun‐exposed locations within them) and residual tree injuries caused by harvesting operations. These conditions attracted beetles such as Trypodendron lineatum (Scolytinae) and Rhagium inquisitor (Cerambycidae), two species known for their ability to attack weakened, dying and dead hosts. Most species were more abundant in selection cuts, except for Evodinus m. monticola (Cerambycidae) whose abundance was strongly reduced after treatment. Some beetles can have detrimental effects on residual trees and thus could reduce timber value, but most species found in treated stands do not represent a high risk for healthy trees. Thus, selection cuts do not seem favourable to the establishment of tree‐killing beetles. However, as they were found more active/abundant after selection cutting, it would be wise to further study their population dynamics over mid‐ and long‐term periods, along with the ecological and economic implications associated with this silvicultural treatment.     

A global change-induced biome shift in the Montseny mountains (NE Spain)

Shifts in plant species and biome distribution in response to warming have been described in past climate changes. However, reported evidence of such shifts under current climate change is still scarce. By comparing current and 1945 vegetation distribution in the Montseny mountains (Catalonia, NE Spain), we report here a progressive replacement of cold‐temperate ecosystems by Mediterranean ecosystems. Beech (Fagus sylvatica) forest has shifted altitudinally upwards by ca. 70 m at the highest altitudes (1600–1700 m). Both the beech forests and the heather (Calluna vulgaris) heathlands are being replaced by holm oak (Quercus ilex) forest at medium altitudes (800–1400 m). This beech replacement has been observed to occur through a progressive isolation and degradation of beech stands. In ‘isolated’ (small and surrounded by holm oaks) beech stands, beech trees are 30% more defoliated, beech recruitment is 41% lower, and holm oak recruitment is three times higher than in ‘continental’ (large and continuous) beech stands. The progressively warmer conditions, complemented by the land use changes (mainly the cessation of traditional land management) are the apparent causes, providing a paradigmatic example of global change affecting distributions of plant species and biomes.     

Spruce beetle outbreak was not driven by drought stress: Evidence from a tree‐ring iso‐demographic approach indicates temperatures were more important

Climate change has amplified eruptive bark beetle outbreaks over recent decades, including spruce beetle (Dendroctonus rufipennis). However, for projecting future bark beetle dynamics there is a critical lack of evidence to differentiate how outbreaks have been promoted by direct effects of warmer temperatures on beetle life cycles versus indirect effects of drought on host susceptibility. To diagnose whether drought‐induced host‐weakening was important to beetle attack success we used an iso‐demographic approach in Engelmann spruce (Picea engelmannii) forests that experienced widespread mortality caused by spruce beetle outbreaks in the 1990s, during a prolonged drought across the central and southern Rocky Mountain region. We determined tree death date demography during this outbreak to differentiate early‐ and late‐dying trees in stands distributed across a landscape within this larger regional mortality event. To directly test for a role of drought stress during outbreak initiation we determined whether early‐dying trees had greater sensitivity of tree‐ring carbon isotope discrimination (?¹³C) to drought compared to late‐dying trees. Rather, evidence indicated the abundance and size of host trees may have modified ?¹³C responses to drought. ?¹³C sensitivity to drought did not differ among early‐ versus late‐dying trees, which runs contrary to previously proposed links between spruce beetle outbreaks and drought. Overall, our results provide strong support for the view that irruptive spruce beetle outbreaks across North America have primarily been driven by warming‐amplified beetle life cycles whereas drought‐weakened host defenses appear to have been a distant secondary driver of these major disturbance events.     

Robinia pseudoacacia as a surrogate for native tree species for saproxylic beetles inhabiting the riparian mixed forests of northern Italy

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A Technique to Screen American Beech for Resistance to the Beech Scale Insect (Cryptococcus fagisuga Lind.)

Beech bark disease (BBD) results in high levels of initial mortality, leaving behind survivor trees that are greatly weakened and deformed. The disease is initiated by feeding activities of the invasive beech scale insect, Cryptococcus fagisuga, which creates entry points for infection by one of the Neonectria species of fungus. Without scale infestation, there is little opportunity for fungal infection. Using scale eggs to artificially infest healthy trees in heavily BBD impacted stands demonstrated that these trees were resistant to the scale insect portion of the disease complex¹. Here we present a protocol that we have developed, based on the artificial infestation technique by Houston², which can be used to screen for scale-resistant trees in the field and in smaller potted seedlings and grafts. The identification of scale-resistant trees is an important component of management of BBD through tree improvement programs and silvicultural manipulation.     

Drought and stand susceptibility to attacks by the European spruce bark beetle: A remote sensing approach

1. Several time-series analyses have demonstrated that after extreme summer drought bark beetle damage increased. However, studies predicting stand susceptibility over large spatial extents are limited by technical constraints in obtaining detailed, spatially-explicit data on infestation spot occurrence.
2. Using a unique dataset of georeferenced bark beetle infestation data, we tested whether the spatial variation of local growing conditions of forest stands, topography, and landscape variables modified the local occurrence of Ips typographus infestations after a severe hot drought in Central Europe.
3. Bark beetle infestation occurrence depended on soil-related aridity intensity, elevation, slope, and soil conditions. We showed that elevation interacted with growing conditions and topography. At low elevations, spruce forests growing on flat areas and wetter soils were more sensitive to the infestations. On the contrary, forests on steep slopes and soils with low water availability were rarely attacked. At the landscape scale, bark beetle damage increased with host tree cover but decreased with compositional diversity.
4. Our findings are generally consistent with the growth-differentiation balance hypothesis that predicts that trees growing under chronic dry conditions tend to be more resistant against biotic disturbances.
5. Spruce stands at low elevations located in homogeneous landscapes dominated by spruce were those more exposed to bark beetles in the initial phase of a drought-induced outbreak.

     

Sources of variation in the incidence of ant–aphid mutualism in boreal forests

• 1 The mutualism between wood ants of the Formica rufa group and aphids living in the canopy of trees is a widespread phenomenon in boreal forests, and it can affect tree growth. However, not all trees in the forest are involved in this interaction.
• 2 To assess the incidence of host trees involved in this ant–aphid mutualism and its spatial distribution in boreal forests, we inventoried sample plots with a radius of 10–15 m around wood ant mounds in 12 forest stands of two age classes (5–12‐year‐old sapling stands and 30–45‐year‐old pole stands) and two dominant tree species (Scots pine and silver birch) in Eastern Finland from 2007 to 2009.
• 3 The proportion of trees visited by ants out of all trees on the individual study plots were in the range 4–62%, and 1.5–39% of the trees on the plots were consistently visited by ants during all 3 years. The percentage of host trees increased with the ant mound base area on the plots. Trees visited by ants were larger and closer to the mound than trees not visited by ants. Within the group of visited trees, more ants were found on bigger trees and on trees close to the ant mounds.
• 4 Extrapolated from plot to stand level, we estimated that 0.5–6.6% of the trees were host trees in at least one of the three study years, and that only 0.01–2.3% of all the trees were consistently visited by ants during all 3 years. It is concluded that ant–aphid mutualism is a minor occurrence at the stand level.

     

Population dynamics of the beech caterpillar, <Emphasis Type="Italic">Syntypistis punctatella</Emphasis>, and biotic and abiotic factors

The beech caterpillar, Syntypistis punctatella (Motschulsky) (Lepidoptera: Notodontidae), often causes extensive defoliation of beech forests in Japan. Outbreaks have often occurred synchronously among different areas at intervals of 8–11 years. Synchrony of outbreaks was considered to be caused by synchrony of weather. Populations of this insect exhibit periodical dynamics in both outbreak and nonoutbreak areas. Factors that might influence the population dynamics of the beech caterpillar were classified from the point of view of the natural bioregulation com-plex, which includes a coleopteran predator, Calosoma maximowiczi, avian predators, parasitoids, entomopathogenic fungi, and delayed induced defensive response (DIR) of beech trees. Because such periodic population dynamics are believed to be caused by one or more delayed density-dependent factors, delayed density-dependent mortality has been identified as a likely source of population cycles. The DIR and pathogenic diseases showed a high order of density dependence. An infectious pathogen, Cordyceps militaris, was considered to be the most plausible agent responsible for periodic dynamics of the beech caterpillar population because insect diseases were effective in cases in which the S. punctatella population started to decrease without reaching outbreak densities, but DIR was not. Conspicuous defoliation caused by this insect tends to occur at certain elevations, where forests are composed of pure stands of beech trees. I propose three different hypotheses to explain this phenomenon: the diversity–stability hypothesis, the resource concentration hypothesis, and the altitudinal soil nutrient hypothesis. Received: November 20, 1999 / Accepted: August 3, 2000     

Prescribed fire is associated with increased floral richness and promotes short-term increases in bee biodiversity in the ponderosa pine forest of the Southern Rocky Mountains

1. Managed low-severity surface fires are frequently implemented in efforts to restore disturbance processes to forests of North America; although the effects of managed fire on forest structure are well-studied, few studies investigate whether these disturbances cascade to impact pollinator communities.
2. We analysed bee-habitat relationships in fire-treated (1- and 3-years post-treatment) and non-treated ponderosa pine stands in Colorado to test wild bee population responses.
3. Observed bee richness and α-diversity were highest in stands 1-year post-fire and had more Anthophora, Bombus, Osmia and Lasioglossum spp. in comparison to 3-year post-fire and non-treated stands. Bee functional groups were responsive to treatments, with more below-ground nesting taxa present in stands 3 years post-fire.
4. Floral richness was the highest mid-growing season (June, July) and within 1-year post-fire stands.
5. A model analysing the effects of foraging and nesting habitat variation on bee assemblages indicated positive association between floral richness and bee α-diversity, but negative relationships with stand basal area. Nesting habitat was not associated with variation in bee assemblages.
6. We conclude that managed fire has positive short-term effects on bee biodiversity that are likely mediated by floral richness. However, these effects were not detectable by 3 years post-treatment in the southern Rocky Mountain region.

     

Changes in diversity and storage function of ectomycorrhiza and soil organoprofile dynamics after introduction of beech into Scots pine forests

Diversity and storage function of mycorrhiza as well as soil organoprofile formation were investigated in a chronosequence of a pure Scots pine (Pinus sylvestris L.) stand, of Scots pine stands that were underplanted with beech (Fagus sylvatica L.) and in three pure beech stands of different age. Mycorrhiza diversity was higher in the pure beech stands compared to the pure pine stand. Beech and pine trees in the mixed stands had similar dominant mycorrhiza morphotypes. However, trees in two of the three pure beech stands were mycorrhized with other types. Mycorrhizal abundance and nutrient amounts of mycorrhizae associated with beech trees were higher in the mixed and in the pure beech stands compared to pine mycorrhizae indicating that nutrient uptake was higher in older beech than in older pine trees. Humus quality varied from pine to beech stands. Plant litter storage in the humus layer was highest in the youngest mixed stand and lowest in the oldest beech stand. Humus forms changed from moder grass-type in the pure Scots pine stand to mor-like moder and moder rich in fine humus with increasing age of beeches in the mixed stands. The older beech stands were characterised by oligomull and mull-like moder as the dominating humus forms. The ecologically favourable humus forms, i.e., nutrient rich humus forms in the older beech stands correlate well with the higher mycorrhizal diversity and abundance as well as the higher nutrient storage of their mycorrhizae in these stands. The results are also discussed with regard to the 'base-pump effect' of beech trees.     

Phenology and growth of Fagus sylvatica and Quercus robur seedlings in response to temperature variation in the parental versus offspring generation

• Plants are known to respond to warming temperatures. Few studies, however, have included the temperature experienced by the parent plant in the experimental design, in spite of the importance of this factor for population dynamics.
• We investigated the phenological and growth responses of seedlings of two key temperate tree species (Fagus sylvatica and Quercus robur) to spatiotemporal temperature variation during the reproductive period (parental generation) and experimental warming of the offspring. To this end, we sampled oak and beech seedlings of different ages (1–5 years) from isolated mother trees and planted the seedlings in a common garden.
• Warming of the seedlings advanced bud burst in both species. In oak seedlings, higher temperatures experienced by mother trees during the reproductive period delayed bud burst in control conditions, but advanced bud burst in heated seedlings. In beech seedlings, bud burst timing advanced both with increasing temperatures during the reproductive period of the parents and with experimental warming of the seedlings. Relative diameter growth was enhanced in control oak seedlings but decreased with warming when the mother plant experienced higher temperatures during the reproductive period.
• Overall, oak displayed more plastic responses to temperatures than beech. Our results emphasise that temperature during the reproductive period can be a potential determinant of tree responses to climate change.