Spatial analysis of gypsy moth populations in Sardinia using geostatistical and climate models

• 1 Spatial fluctuations of the Sardinian population of the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae) were characterized using geostatistical and climate models. Data on gypsy moth egg mass abundance recorded at 282 permanent monitoring sites from 1980 to 2004 were incorporated in a geographic information system with the vegetational, geomorphological and pedological features of the sites.
• 2 Statistical analyses revealed that the relative outbreak frequency was related to the predominant host tree, slope and elevation of the monitoring sites, whereas there was no correlation between outbreak frequency and exposure and soil type.
• 3 By using bioclimatic modelling, probability maps of gypsy moth outbreaks were generated. The model identified a probability surface with climatic conditions favourable to gypsy moth outbreaks and thus potentially subject to defoliation. The maps included 92 sites where outbreaks never occurred, suggesting that the Sardinian climate may not be a determinant factor for gypsy moth outbreaks.
• 4 The geostatistical method cokriging with outbreak frequency as a covariate was found to be the most suitable technique to estimate gypsy moth egg mass abundance. Semivariograms showed spatial correlation of egg mass abundance within the range 18.5–53 km. The results obtained were used to create regional gypsy moth distribution maps by cokriging, which demonstrated the outbreak foci and different infestation levels at each monitoring area. These results can help to delimit the treatment areas and develop rational gypsy moth management programmes.

     

Distribution of the entomopathogenic fungus Entomophaga maimaiga (Entomophthorales: Entomophthoraceae) at the northern edge of its range in Europe

The gypsy moth Lymantria dispar is a serious economic pest in European broadleaf forests. However, the entomopathogenic fungus Entomophaga maimaiga, which has a great potential to regulate gypsy moth numbers, has recently spread in the Central and Eastern European area of the moth's range. In the current study, 39 plots in oak forests in the Slovak Republic and the Czech Republic were monitored for E. maimaiga from 2014 to 2016. These plots were located along the northern edge of the E. maimaiga range where gypsy moth outbreaks have occurred in the past. The fungus was detected in 16 of the 39 plots. The results thus confirm that E. maimaiga is quite widespread along the northern edge of its range in Europe and can be considered to be established in that area.     

Extensive gypsy moth defoliation in Southern New England characterized using Landsat satellite observations

Southern New England is currently experiencing the first major gypsy moth (Lymantria dispar) defoliation event in nearly 30 years. Using a novel approach based on time series of Landsat satellite observations, we generated consistent maps of gypsy moth defoliation for 2015 (first year of the outbreak), 2016 (second year of outbreak), and 2017 (third year of outbreak). Our mapped results demonstrate that the defoliation event continued through the 2017 growing season. Moreover, the affected area more than doubled in extent each year and expanded radially to encompass 4386 km² of forested area in Rhode Island, eastern Connecticut, and central Massachusetts. The current gypsy moth outbreak is believed to be the result of a series of unusually dry springs in 2014, 2015, and 2016, which suppressed Entomophaga maimaiga, a fungal mortality agent that has historically reduced gypsy moth impacts in this region. The continuation and marked expansion of the outbreak in 2017 despite average spring rainfall suggests that caterpillars were active early in the growing season, and mortality from the fungus likely peaked after significant defoliation had already occurred. Our Landsat time series approach represents an important new source of data on spatial and temporal patterns in gypsy moth defoliation, and continued satellite-based monitoring will be essential for tracking the progress of this and other gypsy moth outbreaks.     

Climate change and the outbreak ranges of two North American bark beetles

Abstract

• 1 One expected effect of global climate change on insect populations is a shift in geographical distributions toward higher latitudes and higher elevations. Southern pine beetle Dendroctonus frontalis and mountain pine beetle Dendroctonus ponderosae undergo regional outbreaks that result in large‐scale disturbances to pine forests in the south‐eastern and western United States, respectively.
• 2 Our objective was to investigate potential range shifts under climate change of outbreak areas for both bark beetle species and the areas of occurrence of the forest types susceptible to them.
• 3 To project range changes, we used discriminant function models that incorporated climatic variables. Models to project bark beetle ranges employed changed forest distributions as well as changes in climatic variables.
• 4 Projected outbreak areas for southern pine beetle increased with higher temperatures and generally shifted northward, as did the distributions of the southern pine forests.
• 5 Projected outbreak areas for mountain pine beetle decreased with increasing temperature and shifted toward higher elevation. That trend was mirrored in the projected distributions of pine forests in the region of the western U.S. encompassed by the study.
• 6 Projected outbreak areas for the two bark beetle species and the area of occurrence of western pine forests increased with more precipitation and decreased with less precipitation, whereas the area of occurrence of southern pine forests decreased slightly with increasing precipitation.
• 7 Predicted shifts of outbreak ranges for both bark beetle species followed general expectations for the effects of global climate change and reflected the underlying long‐term distributional shifts of their host forests.

     

Ecosystem Impacts of a Range Expanding Forest Defoliator at the Forest-Tundra Ecotone

Insect outbreaks in northern-boreal forests are expected to intensify owing to climate warming, but our understanding of direct and cascading impacts of insect outbreaks on forest ecosystem functioning is deficient. The duration and severity of outbreaks by geometrid moths in northern Fennoscandian mountain birch forests have been shown to be accentuated by a recent climate-mediated range expansion, in particular of winter moth (Operophtera brumata). Here, we assess the effect of moth outbreak severity, quantified from satellite-based defoliation maps, on the state of understory vegetation and the abundance of key vertebrate herbivores in mountain birch forest in northern Norway. We show that the most recent moth outbreak caused a regional-scale state change to the understory vegetation, mainly due to a shift in dominance from the allelopathic and unpalatable dwarf-shrub Empetrum nigrum to the productive and palatable grass Avenella flexuosa. Both these central understory plant species responded significantly and nonlinearly to increasing outbreak severity. We further provide evidence that the effects of the outbreak on understory vegetation cascaded to cause strong but opposite impacts on the abundance of the two most common herbivore groups. Rodents increased with defoliation, largely mirroring the increase in A. flexuosa, whereas ungulate abundance instead showed a decreasing trend. Our analyses also suggest that the response of understory vegetation to defoliation may depend on the initial state of the forest, with poorer forest types potentially allowing stronger responses to defoliation.     

Spatial analysis of harmonic oscillation of gypsy moth outbreak intensity

Outbreaks of many forest-defoliating insects are synchronous over broad geographic areas and occur with a period of approximately 10 years. Within the range of the gypsy moth in North America, however, there is considerable geographic heterogeneity in strength of periodicity and the frequency of outbreaks. Furthermore, gypsy moth outbreaks exhibit two significant periodicities: a dominant period of 8–10 years and a subdominant period of 4–5 years. In this study, we used a simulation model and spatially referenced time series of outbreak intensity data from the Northeastern United States to show that the bimodal periodicity in the intensity of gypsy moth outbreaks is largely a result of harmonic oscillations in gypsy moth abundance at and above a 4 km² scale of resolution. We also used geographically weighted regression models to explore the effects of gypsy moth host-tree abundance on the periodicity of gypsy moths. We found that the strength of 5-year cycles increased relative to the strength of 10-year cycles with increasing host tree abundance. We suggest that this pattern emerges because high host-tree availability enhances the growth rates of gypsy moth populations.     

Stand‐replacing fires reduce susceptibility of lodgepole pine to mountain pine beetle outbreaks in Colorado

Aim As climate change is increasing the frequency, severity and extent of wildfire and bark beetle outbreaks, it is important to understand how these disturbances interact to affect ecological patterns and processes, including susceptibility to subsequent disturbances. Stand‐replacing fires and outbreaks of mountain pine beetle (MPB), Dendroctonus ponderosae, are both important disturbances in the lodgepole pine, Pinus contorta, forests of the Rocky Mountains. In the current study we investigated how time since the last stand‐replacing fire affects the susceptibility of the stand to MPB outbreaks in these forests. We hypothesized that at a stand‐scale, young post‐fire stands (< c. 100–150 years old) are less susceptible to past and current MPB outbreaks than are older stands. Location Colorado, USA. Methods We used dendroecological methods to reconstruct stand‐origin dates and the history of outbreaks in 23 lodgepole pine stands. Results The relatively narrow range of establishment dates among the oldest trees in most sampled stands suggested that these stands originated after stand‐replacing or partially stand‐replacing fires over the past three centuries. Stands were affected by MPB outbreaks in the 1940s/1950s, 1980s and 2000s/2010s. Susceptibility to outbreaks generally increased with stand age (i.e. time since the last stand‐replacing fire). However, this reduced susceptibility of younger post‐fire stands was most pronounced for the 1940s/1950s outbreak, less so for the 1980s outbreak, and did not hold true for the 2000s/2010s outbreak. Main conclusions Younger stands may not have been less susceptible to the most recent outbreak because: (1) after stands reach a threshold age of > 100–150 years, stand age does not affect susceptibility to outbreaks, or (2) the high intensity of the most recent outbreak reduces the importance of pre‐disturbance conditions for susceptibility to disturbance. If the warm and dry conditions that contribute to MPB outbreaks concurrently increase the frequency and/or extent of severe fires, they may thereby mitigate the otherwise increased landscape‐scale susceptibility to outbreaks. Potential increases in severe fires driven by warm and dry climatic trends may lead to a negative feedback by making lodgepole pine stands less susceptible to future MPB outbreaks.     

Life history and behavioural traits of Mnesampela privata that exacerbate population responses to eucalypt plantations: Comparisons with Australian and outbreak species of forest geometrid from the Northern Hemisphere

The larvae of the native Australian moth Mnesampela privata (Geometridae) sometimes defoliate plantation eucalypts, causing concern to industry. Typically, populations of M. privata maintain innocuous numbers in native forests, but outbreak populations can occur in plantations. Certain of the life history and behavioural traits of M. privata exacerbate population responses in simplified systems. Specifically, M. privata exhibits indiscriminate oviposition behaviour, for example, females oviposit upon leaves where conspecific egg clutches are already present. Combined with large egg clutches, this trait can lead to heavy exploitation of natal hosts. Complete defoliation of the natal tree necessitates that larvae disperse to undamaged hosts. In native forests, dispersing larvae would have a lower probability of locating a new host of suitable species and phenotype than they would in a plantation. Larval tolerance of a wide variety of species of Eucalyptus facilitates utilization of non‐natal eucalypts. A lower rate of encounter between larvae and natural enemies is achieved by means of leaf shelters and autumnal seasonal activity. Good dispersion by gravid females from plantations with large moth populations ensures that nearby plantations will be located. Given these characteristics, continued outbreaks of M. privata are likely in monospecific eucalypt plantations, especially when management practices reduce populations of natural enemies and overall vegetational complexity. However, because outbreaks only arise in plantations, population fluctuations of M. privata cannot be considered ‘self‐driven’. Hence, this moth is not a true eruptive species in the manner of other geometrids such as the autumnal moth (Epirrita autumnata autumnata) from Europe or the hemlock looper (Lambdina fiscellaria fiscellaria) from North America, but rather a gradient outbreak insect.     

Spatial and temporal dynamics of Siberian silk moth large-scale outbreak in dark-needle coniferous tree stands in Altai

The spatial and temporal dynamics of fir stands damage caused by a large-scale outbreak of the Siberian silk moth (Dendrolimus sibiricus Tschetv.) in the Altai Mountains has been studied using remote sensing and GIS methods. It is ascertained that forest damages are dissimilar relative to the surface features. The pest outbreak initially occurred on the southwest slopes with mean values of 10° and elevation of 400 m asl. The damages further extended both upward and downward, involving slopes of high steepness and the eastern exposure. The total area of the dead stands comprised 6000 ha, 45% of which were lost due to secondary pest (xylophagous insects) attacks. It is indicated that the use of remote sensing made it possible to determine the beginning of a pest outbreak with one-decade precision. The Siberian silk moth large-scale outbreak occurred against an increase in air temperatures, a decrease in precipitation, and a reduction in late frosts. The tree plants weakened by the Siberian silk moth and water stress were affected by xylophagous insect attacks. The observed and predicted warming and climate aridity increase will facilitate Siberian silk moth outbreaks both within its range and northward of the current margins of the range.     

Large-scale forest fragmentation increases the duration of tent caterpillar outbreak

I examined historical data (1950–1984) on the duration of outbreaks of the forest tent caterpillar (Malacosoma disstria) in northern Ontario, Canada. Outbreak duration was compared to host tree species dominance and forest structure over large areas of boreal forest partially cleared for agriculture. Abundance of the principal host tree species Populus tremuloides had no consistent effect on duration of outbreak within forest districts, and was negatively correlated with duration of outbreaks among the eight forest districts examined. The amount of forest edge per km² was the best, and most consistent, predictor of the duration of tent caterpillar outbreaks both within individual forest districts and among forest districts. Because forest tent caterpillar populations are driven largely by the impact of parasitoids and pathogens, results here suggest that large-scale increase in forest fragmentation affects the interaction between these natural enemies and forest tent caterpillar. Increased clearing and fragmentation of boreal forests, by agriculture and forestry, may be exacerbating outbreaks of this forest defoliator.     

Geographical variation in the periodicity of gypsy moth outbreaks

The existence of periodic oscillations in populations of forest Lepidoptera is well known. While information exists on how the periods of oscillations vary among different species, there is little prior evidence of variation in periodicity within the range of a single Lepidopteran species. The exotic gypsy moth is an introduced foliage-feeding insect in North America. Its populations are characterized by high amplitude oscillations between innocuously low densities and outbreak levels during which large regions of forests are defoliated. These outbreaks are recognized to arise periodically with considerable synchrony across much of the gypsy moth's North American range. Our analysis indicates that gypsy moth outbreaks in North America are periodic but they exhibit two dominant periodicities: a primary period of 8–10 yr (as previously reported) and a secondary period of 4–5 yr (a new finding in this study). The outbreak periodicity varied geographically and this variation was associated with forest type. We found that in the most susceptible forest types, those on xeric sites where oak is often mixed with pines, outbreak periodicity had a more dominant 5-yr period while in forest types characteristic of more mesic sites where oak was mixed with maples and other species, cycles were clearly operating on a 10-yr period.     

Combined Effects of Energy Development and Disease on Greater Sage-Grouse

Species of conservation concern are increasingly threatened by multiple, anthropogenic stressors which are outside their evolutionary experience. Greater sage-grouse are highly susceptible to the impacts of two such stressors: oil and gas (energy) development and West Nile virus (WNv). However, the combined effects of these stressors and their potential interactions have not been quantified. We used lek (breeding ground) counts across a landscape encompassing extensive local and regional variation in the intensity of energy development to quantify the effects of energy development on lek counts, in years with widespread WNv outbreaks and in years without widespread outbreaks. We then predicted the effects of well density and WNv outbreak years on sage-grouse in northeast Wyoming. Absent an outbreak year, drilling an undeveloped landscape to a high permitting level (3.1 wells/km²) resulted in a 61% reduction in the total number of males counted in northeast Wyoming (total count). This was similar in magnitude to the 55% total count reduction that resulted from an outbreak year alone. However, energy-associated reductions in the total count resulted from a decrease in the mean count at active leks, whereas outbreak-associated reductions resulted from a near doubling of the lek inactivity rate (proportion of leks with a last count = 0). Lek inactivity quadrupled when 3.1 wells/km² was combined with an outbreak year, compared to no energy development and no outbreak. Conservation measures should maintain sagebrush landscapes large and intact enough so that leks are not chronically reduced in size due to energy development, and therefore vulnerable to becoming inactive due to additional stressors.     

Outbreaks of the winter moth on Sitka Spruce in Scotland are not influenced by nutrient deficiencies of trees,tree budburst,or pupal predation

Summary Since the early 1980s, the winter moth, Operophtera brumata L. (Lepidoptera: Geometridae) has emerged as a serious pest of Sitka Spruce, Picea sitchensis Bong. plantations in southern Scotland. Outbreaks are characterised by susceptible sites within plantations which can occur immediately adjacent to resistant sites. We investigated the level of some nutrients in the trees, the date of budburst of the trees, and the numbers of some potential predators of winter moth pupae. None could satisfactorily explain outbreak patterns. Although foliage analysis demonstrated that many trees were marginal or deficient in phosphorus, nitrogen and potassium, these deficiencies were not related to the susceptibility of a site. Within sites, the numbers and weights of O. brumata were positively related to phosphorus content and negatively related to calcium content of foliage. Other evidence suggests, however, that these correlations may not represent direct effects of phosphorus and calcium on larval growth and survival. Date of budburst, which commonly determines susceptibility of deciduous hosts to O. brumata, was unrelated to density, and pupal predators were more, not less, abundant in susceptible sites. Although it is difficult to distinguish between factors that initiate outbreaks and those that maintain them, these data suggest that nutrient deficiencies of trees, budburst date, and the distribution of pupal predators of the winter moth cannot explain patterns of outbreak of the winter moth on spruce.     

Population densities and geographic distribution of night monkeys (Aotus nancymai and Aotus vociferans) (cebidae: Primates) in Northeastern Peru

Population densities of two species of night (or owl) monkeys (Aotus nancymai and Aotus vociferans) were estimated using transect census methods. Densities of Aotus nancymai were approximately 46.3 individuals/km² in lowland forests and 24.2 individuals/km² in highland forests. For Aotus vociferans densities were 33.0 individuals/km² in lowland forests and 7.9 individuals/km² in highland forests. A. vociferans occurs north of the Río Amazonas-Marañon and A. nancymai south of it, except for the northern enclave between the Ríos Tigre and Pastaza. The two species are nowhere sympatric. However, the two known karyotypic forms of A. vociferans occupy the same habitats throughout the Peruvian range of the species.     

A simple Bayesian network to interpret the accuracy of armyworm outbreak forecasts

Forecasting of outbreaks of armyworm (larvae of the moth Spodoptera exempta) employs information from rain gauges and moth traps. Rainfall is an independent variable, but moth catch is affected by rainfall, and outbreak risk is affected by both moth catch and rainfall. A simple Bayesian network was used to describe these relationships and so derive conditional probabilities. The data were from a new initiative, community‐based forecasting of armyworm in Tanzania, in which outbreak risk for a village is determined locally from a single moth trap and rain gauge located within the village. It was found that, following a positive forecast, an armyworm outbreak was approximately twice as likely to occur as would be expected by chance. If the forecast was negative because of insufficient moths, outbreaks were half as likely as would be expected by chance. If the forecast was negative because of insufficient rain, however, the outbreak probability remained similar to chance: an aspect of the forecast that requires improvement. Overall, a high forecasting accuracy can be achieved by village communities using simple rules to predict armyworm outbreaks.     

Shifting altitudinal distribution of outbreak zones of winter moth Operophtera brumata in sub‐arctic birch forest: a response to recent climate warming?

Climatic change is expected to affect the extent and severity of geometrid moth outbreaks, a major disturbance factor in sub-arctic birch forests. Previous studies have reported that the two geometrid species involved, autumnal moth and winter moth, differ in their temperature requirements and, consequently, in their altitudinal and latitudinal distribution patterns. In this study, we document the altitudinal distribution of winter moth outbreaks in a large coastal area in northern Norway. We show that, in the present winter moth outbreak, defoliated birch stands were seen as distinct zones with a rather constant width in the uppermost part of the forest and where the upper limit coincided with the forest line. The outbreak zone closely followed the spatially variable forest line as an undulating belt, although some of the variation in outbreak zone width was also related to variation in topographical variables, such as distance from the coast, forest line altitude, and slope of the terrain. A distinct outbreak zone at the altitudinal forest line is the typical picture that has been depicted in more qualitative historical records on previous outbreaks of autumnal moth rather than winter moth. We suggest that the recent documented climate warming in this region may have induced a shift in distribution of the winter moth both relative to topography and geography. Further investigation is, however, required to substantiate these suspicions.     

A survey of elephants (Loxodonta africana) in the Kahuzi-Biega National Park lowland sector and adjacent forest in eastern Zaire

There has been considerable uncertainty about the abundance estimation of Loxodonta africana within tropical lowland moist forests in Zaire. We surveyed a 15,570 km² area within the forests of eastern Zaire using transect sampling methods and estimated the elephant population to be 3720 (range 2300–5000) individuals. Dung pile densities were significantly different between adjacent settlement, deep forest, and deep forest core strata, with the most remote area harbouring the highest density. Evidence of elephant poaching was encountered throughout the survey area suggesting that elephant populations continue to be at severe risk.     

Growth and survival of larvae of Thaumetopoea pinivora inside and outside a local outbreak area

1 Outbreaks of herbivorous insects tend to be spatially restricted, possibly because of demographic differences between inside and outside the outbreak area. In some cases, the margin of the outbreak area is distinct, allowing comparisons of adjacent areas that may identify factors leading to such differences in abundance. The northern pine processionary moth Thaumetopoea pinivora presently occurs at outbreak densities within a well‐defined area of approximately 3000 ha on the island of Gotland, south Sweden. We investigated whether cohorts of young larvae (first and second instar) had higher growth rate and survival inside or outside the outbreak area. 2 Group‐feeding appears to promote outbreaks in certain insect groups. Because T. pinivora larvae are highly social, we also compared larval performance between groups of different sizes inside and outside of the outbreak area: ‘small’ (33 eggs/group) and ‘normal’ (100 eggs/group). 3 Averaged over group size, whole colony mortality through the first two instars was two‐fold higher in the non‐outbreak area compared with the outbreak area. Mortality of individual larvae in the surviving colonies, however, did not differ between the two areas. There were only small differences in food quality (toughness, nitrogen content) between the areas, with no detectable effects on larval performance. 4 Larval relative growth rate did not differ between reduced and normal‐sized groups, which is surprising given that growth rate is known to increase with group size in other group‐feeding lepidopterans. 5 Reduced group size negatively affected larval survival, particularly in the outbreak area; by contrast, normal‐sized groups survived equally well in the two areas. Wood ants (Formica spp.) were more common outside the outbreak area, and appeared to be the main cause of colony mortality at low larval density. A different result was observed with regard to per‐capita mortality, which was higher in the outbreak area. We speculate that this could have been due to solitary predators being locally specialized on T. pinivora in the high‐density area.     

Sleeping Parties and Nest Distribution of Chimpanzees in the Savanna Woodland, Ugalla, Tanzania

We conducted ecological studies of chimpanzees (Pan troglodytes) in the Ugalla area, western Tanzania. Ugalla is one of the driest habitats of chimpanzees and the Ugalla River is the eastern boundary of chimpanzee distribution. Most of Ugalla is occupied by savanna woodlands dominated by deciduous trees of Brachystegia and Julbernardia. Chimpanzees tended not to make nests in riverine forests in plains, but in small patchy forests dominated by Monopetalanthus richardsiae and valley forests dominated by Julbernardia unijugata on slopes in mountainous areas. We estimated population density of chimpanzees to be 7–9 × 10⁻² individuals/km² based on nest censuses, suggesting that 2–3 × 10² individuals inhabited the 3352 km² area of Ugalla. The size of the largest nest cluster (n=23) suggests that 1 unit group (community) comprised 30–35 individuals. In the daytime, chimpanzees formed small feeding parties (mean 2.0 individuals), but larger ones in the evening (mean 4.8 individuals and 5.2 individuals based on fresh nest clusters). The pattern might reduce the predation risk from large nocturnal carnivores such as lions and leopards. The sleeping sites may function as both a safe sleeping site and a meeting point for chimpanzees with a huge home range that may have difficulty in finding other members of their unit group.     

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.