Drought then wildfire reveals a compound disturbance in a resprouting forest.

The frequency and intensity of forest disturbances, such as drought and fire, are increasing globally, with an increased likelihood of multiple disturbance events occurring in short succession. Disturbances layered over one another may influence the likelihood or intensity of subsequent events (a linked disturbance) or impact response and recovery trajectories (a compound disturbance), with substantial implications for ecological spatiotemporal vulnerability. This study evaluates evidence for disturbance interactions of drought followed by wildfire in a resprouting eucalypt-dominated forest (the Northern Jarrah Forest) in southwestern Australia. Sites were stratified by drought (high, low), from previous modeling and ground validation, and fire severity (high, moderate, unburnt), via remote sensing using the relative difference normalized burn ratio (RdNBR). Evidence of a linked disturbance was assessed via fine fuel consumption and fire severity. Compound disturbance effects were quantified at stand scale (canopy height, quadratic mean diameter, stem density) and stem scale (mortality). There was no evidence of prior drought influencing fine fuel consumption or fire severity and, hence, no evidence of a linked disturbance. However, compound disturbance effects were evident; stands previously affected by drought experienced smaller shifts in canopy height, quadratic mean diameter, and stem density than stands without prior drought impact. At the stem scale, size and fire severity were the strongest determinants of stem survival. Proportional resprouting height was greater in high drought sites than in low drought sites (p < 0.01), meaning, structurally, the low drought stands decreased in height more than the high drought stands. Thus, a legacy of the drought was evident after the wildfire. Although these resprouting eucalypt forests have been regarded as particularly resilient, this study illustrates how multiple disturbances can overwhelm the larger tree component and promote an abundance of smaller stems. We suggest that this is early evidence of a structural destabilization of these forests under a more fire-prone, hotter, and drier future climate.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Within-tree Distribution and Survival of the Eucalyptus Longhorned Borer Phoracantha semipunctata (Coleoptera: Cerambycidae) in a Mediterranean-Type Ecosystem.

The attack patterns, infestation success and larval development of woodborers within living trees are complex and are largely shaped by host tree characteristics. Following a severe drought in a native eucalypt forest where outbreak densities of a native Australian beetle, the eucalyptus longhorned borer (Phoracantha semipunctata), occurred, a tree dissection study was conducted in Australia. This involved felling 40 trees each of jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) that were cut into 1-m sections and neonate larval galleries, larvae in pupal cells and adult borer emergence were measured and added to give total numbers per tree to determine the within-tree distribution and survival of P. semipunctata. There was a significant impact on larval survival in both species, in contrast, pupal survival remained high. Within-tree distribution of P. semipunctata was directional with borer emergence and incidence of larval galleries both negatively associated with tree section height above the ground and positively associated with section diameter and bark thickness, reaching a maximum towards the base of trees. High incidence and survival in lower thicker tree sections indicate a more conducive environment for larval development, in contrast to poor larval survival in smaller thinner sections at the top of trees. The dependence of larval survival on tree characteristics controlling the within-tree distribution of borer emergence is emphasized, and needs to be considered when estimating the spread of borer populations during outbreaks.

Carbon consequences of drought differ in forests that resprout.

Prolonged drought and intense heat-related events trigger sudden forest die-off events and have now been reported from all forested continents. Such die-offs are concerning given that drought and heatwave events are forecast to increase in severity and duration as climate change progresses. Quantifying consequences to carbon dynamics and storage from die-off events are critical for determining the current and future mitigation potential of forests. We took stand measurements five times over 2+ years from affected and unaffected plots across the Northern Jarrah Forest, southwestern Australia, following an acute drought/heatwave in 2011. We found a significant loss of live standing carbon (49.3 t ha-1 ), and subsequently a significant increase in the dead standing carbon pool by 6 months post-die-off. Of the persisting live trees, 38% experienced partial mortality contributing to the rapid regrowth and replenishment (82%-88%) of labile carbon pools (foliage, twigs, and branches) within 26 months. Such regrowth was not substantial in terms of net carbon changes within the timeframe of the study but does reflect the resprouting resilience of this forest type. Dead carbon generated by the die-off may persist for centuries given low fragmentation and decay rates resulting in low biogenic emission rates relative to other forest types. However, future fire may threaten persistence of both dead and live pools via combustion and mortality of live tissue and impaired regrowth capacity. Resprouting forests are commonly regarded as resilient systems, however, a changing climate could see vulnerable portions of forests become carbon sources rather than carbon sinks.

Subcontinental heat wave triggers terrestrial and marine, multi-taxa responses.

Heat waves have profoundly impacted biota globally over the past decade, especially where their ecological impacts are rapid, diverse, and broad-scale. Although usually considered in isolation for either terrestrial or marine ecosystems, heat waves can straddle ecosystems of both types at subcontinental scales, potentially impacting larger areas and taxonomic breadth than previously envisioned. Using climatic and multi-species demographic data collected in Western Australia, we show that a massive heat wave event straddling terrestrial and maritime ecosystems triggered abrupt, synchronous, and multi-trophic ecological disruptions, including mortality, demographic shifts and altered species distributions. Tree die-off and coral bleaching occurred concurrently in response to the heat wave, and were accompanied by terrestrial plant mortality, seagrass and kelp loss, population crash of an endangered terrestrial bird species, plummeting breeding success in marine penguins, and outbreaks of terrestrial wood-boring insects. These multiple taxa and trophic-level impacts spanned >300,000 km2-comparable to the size of California-encompassing one terrestrial Global Biodiversity Hotspot and two marine World Heritage Areas. The subcontinental multi-taxa context documented here reveals that terrestrial and marine biotic responses to heat waves do not occur in isolation, implying that the extent of ecological vulnerability to projected increases in heat waves is underestimated.

Spatial Configuration of Drought Disturbance and Forest Gap Creation across Environmental Gradients.

Climate change is increasing the risk of drought to forested ecosystems. Although drought impacts are often anecdotally noted to occur in discrete patches of high canopy mortality, the landscape effects of drought disturbances have received virtually no study. This study characterized the landscape configuration of drought impact patches and investigated the relationships between patch characteristics, as indicators of drought impact intensity, and environmental gradients related to water availability to determine factors influencing drought vulnerability. Drought impact patches were delineated from aerial surveys following an extreme drought in 2011 in southwestern Australia, which led to patchy canopy dieback of the Northern Jarrah Forest, a Mediterranean forest ecosystem. On average, forest gaps produced by drought-induced dieback were moderate in size (6.6 ± 9.7 ha, max = 85.7 ha), compact in shape, and relatively isolated from each other at the scale of several kilometers. However, there was considerable spatial variation in the size, shape, and clustering of forest gaps. Drought impact patches were larger and more densely clustered in xeric areas, with significant relationships observed with topographic wetness index, meteorological variables, and stand height. Drought impact patch clustering was more strongly associated with the environmental factors assessed (R2 = 0.32) than was patch size (R2 = 0.21); variation in patch shape remained largely unexplained (R2 = 0.02). There is evidence that the xeric areas with more intense drought impacts are 'chronic disturbance patches' susceptible to recurrent drought disturbance. The spatial configuration of drought disturbances is likely to influence ecological processes including forest recovery and interacting disturbances such as fire. Regime shifts to an alternate, non-forested ecosystem may occur preferentially in areas with large or clustered drought impact patches. Improved understanding of drought impacts and their patterning in space and time will expand our knowledge of forest ecosystems and landscape processes, informing management of these dynamic systems in an uncertain future.

Comparative behavior of root pathogens in stems and roots of southeastern Pinus species.

Root diseases are expected to become a greater threat to trees in the future due to accidental pathogen introductions and predicted climate changes, thus there is a need for accurate and efficient pathogenicity tests. For many root pathogens, these tests have been conducted in stems instead of roots. It, however, remains unclear whether stem and root inoculations are comparable for most fungal species. In this study we compared the growth and damage caused by five root pathogens (Grosmannia huntii, Grosmannia alacris, Leptographium procerum, Leptographium terebrantis, and Heterobasidion irregulare) in root and stem tissue of two Pinus species by inoculating mature trees and tissue amended agar in the laboratory. Most fungal species tested caused greater damage in roots of both pine hosts following inoculation. The relationship between root and stem damage was, however, similar when most combinations of pathogens were compared. These results suggest that although stem inoculations are not suitable for determining the actual damage potential of a given species, they may be viewed as a useful surrogate for root inoculations when comparing the relative pathogenicity of multiple species. When grown on amended agar, fungal species generally had greater growth in stem tissue, contrasting with the findings from tree inoculations.

Ecology of root-feeding beetles and their associated fungi on longleaf pine in Georgia.

Root-feeding beetles, particularly of the curculionid subfamilies Scolytinae and Molytinae, are known to be effective vectors of Ophiostomatoid fungi. Infestation by these insects and subsequent infection by the Ophiostomatoid fungi may play an important role in accelerating symptom progression in pine declines. To examine the relationship between beetles and fungi in longleaf pine stands, root-feeding curculionids were collected in pitfall traps baited with ethanol and turpentine for 62 wk, and Ophiostomatoid fungi were isolated from their body surfaces. The most abundant root-feeding beetles captured were Hylastes tenuis, H. salebrosus, Pachylobius picivorus, Hylobius pales, and Dendroctonus terebrans. The number of insects captured peaked in spring and fall, although peaks for different insect taxa did not coincide. The most frequently isolated fungi were Grosmannia huntii, Leptographium procerum, L. terebrantis, and L. serpens. Other Ophiostomatoid fungi recovered included Ophiostoma spp. and Pesotum spp. Insect infestation data suggest that Hylastes spp. share an ecological niche, as do Hb. pales and P. picivorus, because the ratios of their fungal symbionts were similar. The fungi associated with D. terebrans suggest that it did not share habitat with the other principle vectors.

Quantitative, nondestructive assessment of beech scale (Hemiptera: Cryptococcidae) density using digital image analysis of wax masses.

Beech scale, Cryptococcus fagisuga Lindinger, is a non-native invasive insect associated with beech bark disease. A quantitative method of measuring viable scale density at the levels of the individual tree and localized bark patches was developed. Bark patches (10 cm(2)) were removed at 0, 1, and 2 m above the ground and at the four cardinal directions from 13 trees in northern New York and 12 trees in northern Michigan. Digital photographs of each patch were made, and the wax mass area was measured from two random 1-cm(2) subsamples on each bark patch using image analysis software. Viable scale insects were counted after removing the wax under a dissecting microscope. Separate regression analyses at the whole tree level for the New York and Michigan sites each showed a strong positive relationship of wax mass area with the number of underlying viable scale insects. The relationships for the New York and Michigan data were not significantly different from each other, and when pooling data from the two sites, there was still a significant positive relationship between wax mass area and the number of scale insects. The relationships between viable scale insects and wax mass area were different at the 0-, 1-, and 2-m sampling heights but do not seem to affect the relationship. This method does not disrupt the insect or its interactions with the host tree.