G. Matusick

Landscape-scale assessment of tree crown dieback following extreme drought and heat in a Mediterranean eucalypt forest ecosystem

Mediterranean regions are under increasing pressure from global climate changes. Many have experienced more frequent extreme weather events such as droughts and heatwaves, which have severe implications for the persistence of forest ecosystems. This study reports on a landscape-scale assessment investigating potential associated factors of crown dieback in dominant tree species following an extreme dry and hot year/summer of 2010/11 in the Northern Jarrah Forest of Western Australia. Analyses focussed on the influence of (i) geology, (ii) topography, (iii) climate, and (iv) fire history. The results showed that trees on specific soils were more likely to show canopy dieback. Generally, trees on rocky soils with low water holding capacity were found to be affected more frequently. Other explanatory factors identified that dieback occurred (i) on sites that were close to rock outcrops, (ii) in areas that received a slightly higher amount of annual rainfall compared to the surrounding landscape, (iii) on sites at high elevations and (vi) on steep slopes, and (v) in areas that were generally slightly warmer than their surroundings. These results expand our understanding of how landscape-scale factors contribute to the effects of an extreme drought and heating event in Mediterranean forest ecosystems, and give indications of where changes are likely to occur within the landscape in the future. The analogues with other Mediterranean climate regions make the results of this study transferable and a starting point for further investigations.

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

ABSTRACT 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, Pachyhbius 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.

Variation in virulence among four root-inhabiting Ophiostomatoid fungi on Pinus taeda L., P. palustris Mill, and P. elliottii Engelm. seedlings

Abstract Ophiostomatoid fungi have been implicated in root disease of pines in the southeastern United States. To understand more about their virulence, inoculation studies were conducted on loblolly (Pinus taeda), longleaf (Pinus palustris), and slash pine (Pinus elliotii). One-year-old bareroot seedlings, after being planted and established for 11 weeks, were wound-inoculated with one of four prominent North American ophiostomatoid fungal species. After three months, a darkened lesion, extending from the point of inoculation was observed for all species. Grosmannia huntii (L. huntii) caused the greatest lesion and occlusion length in loblolly pine and slash pine. Leptographium procerum and L. terebrantis caused similar lesion and occlusion lengths and were the least virulent among the fungi tested. These studies indicate clear virulence differences among the four North American fungi. Grosmannia huntii, previously not known to be pathogenic, was observed causing significant damage compared with other well-known Leptographium species. Finally, lesion and tissue occlusion lengths were significantly smaller in longleaf pine for all fungal species when compared to loblolly and slash pine.

Linking restoration outcomes with mechanism: the role of site preparation, fertilisation and revegetation timing relative to soil density and water content

Global land use and ongoing climate change highlight the importance of ecological restoration as an emerging discipline and underscore the need for successful revegetation techniques. To link mechanistic drivers of seedling establishment with techniques to increase revegetation success, we undertook field-based experiments in degraded peri-urban woodlands in Mediterranean southwestern Australia using two iconic tree species. Over the course of an entire growing season, our objectives were to: (1) characterise soil moisture profiles in relation to site preparation techniques (ripping and created ashbeds) and (2) determine whether early seedling establishment can be increased through site preparation techniques (ripping and ashbeds), plant treatments (various fertilisers and biochar) and early planting during the wet season. Ripping significantly reduced soil compaction and was associated with a significant soil moisture stratification; moisture penetrated to greater depths in soils that were ripped or treated with ashbeds. Particular site preparation techniques (ripping) and early planting significantly increased early establishment, health and growth of tree seedlings. Fertilisation effects varied by species with generally neutral effects on seedlings. Finally, seedlings planted in ripped soils had significantly longer, deeper root systems accessing portions of the soil profile with higher summer moisture. Techniques such as ripping, even in deep sandy soils, reduce soil compaction and alter moisture availability within the soil profile, promoting deeper root growth and thus increasing revegetation success in these degraded Mediterranean ecosystems. Linkage of revegetation outcomes with plant response and physical soil properties associated with particular treatments provides critical knowledge for both restoration scientists and land managers.

The pathogenicity and virulence of four Ophiostomatoid fungi on young Longleaf pine trees

Abstract In southeastern USA, insect vectors transfer pathogenic ophiostomatoid fungi that cause disease in southern pines. During 2007 and 2008, potted longleaf pines (P. palustris Mill.), of similar ages ranging in height from 58 to 198 cm, were inoculated with the following fungi to assess their pathogenicity (and virulence): Grosmannia huntii, Leptographium procerum, L. serpens and L. terebrantis. Seventeen weeks after inoculation, L. terebrantis, L. serpens and G. huntii were found to cause significantly larger lesions and more sapwood discolouration than wounded uninoculated controls. Leptographium terebrantis caused significantly more sapwood discolouration than all other fungi. Despite significant sapwood occlusion after fungal inoculation, no reductions in needle water potentials were observed between treatments. All fungal species were successfully re-isolated from longleaf pine trees.

Quantitative, Nondestructive Assessment of Beech Scale (Hemiptera: Cryptococcidae) Density Using Digital Image Analysis of Wax Masses

ABSTRACT 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 cm2) 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-cm2 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.

Root-Inhabiting Bark Beetles (Coleoptera: Curculionidae) and their Fungal Associates Breeding in Dying Loblolly Pine in Alabama

Root feeding bark beetles in the genus Hylastes Hylastes breeding in loblolly pine across 3 crown (Coleoptera: Curculionidae: Scolytinae) commoncondition classes (healthy, dying, dead). Addition ly carry ophiostomatoid fungi (Ophistomatales: ally, the relationship between ophiostomatoid Ophistomataceae)and collectively contribute to species, their Hylastes species vectors, and tree root disorders of Pinus species around the world condition were investigated. (Jacobs & Wingfield 2001). One of the most damLoblolly pine stands across central Alabama aging root disorders is black-stain root disease have experienced high mortality rates, character of conifers in the Western United States, caused ized by scattered stand mortality with a distinct by the fungal species Leptographium wageneri lack of above-ground pests (Brown & McDowell Kendrick and its primary beetle vector H. nig1968). During an investigation of one stand in rinus (Mannerheim) (Witcosky et al. 1986). ReMay 2008 in central Alabama, dying trees were cently, Hylastes salebrosus Eichhoff and H. tenuis found to be infested with root-inhabiting beetles Eichhoff have been associated with loblolly pine and their associated ophiostomatoid fungi. Six (Pinus taeda L.) decline (Eckhardt et al. 2007), trees were selected based on crown condition, in which is considered an emerging forest health iseluding 2 with green, healthy crowns (healthy), sue in the southeastern United States (Eckhardt 2 with severely chlorotic and thinning crowns et al. 2010). (dying), and 2 with red crowns (dead). Two pri Loblolly pine decline (LPD) is a tree disease mary lateral roots were randomly selected and complex that is characterized by symptoms that excavated to approximately 3 meters from the include thinning tree crowns, growth reductions root collar. Roots were removed from trees and (Eckhardt et al. 2007) and premature mortality in carefully dissected. All insects, including imma localized areas (Brown and McDowell 1968). Eviture stages, were collected from roots and adult dence suggests the complex is the result of interbeetles were identified to species. Root tissue was acting abiotic (Eckhardt & Menard 2008) and biobtained from each brood gallery for the isolation otic (Eckhardt et al. 2007) stress factors. Hylastes of ophiostomatoid fungal species using methods species vector a variety of root-infecting ophiostodescribed in Eckhardt et al. (2007). Following matoid fungi (Klebzig et al. 1991, 1995; Eckhardt surface sterilization, tissue was placed on CSMA et al. 2007; Zanzot et al. 2010) which are patho(malt extract agar containing 800 mg/L of cyclo genic to loblolly pine (Matusick & Eckhardt 2010; heximide and 200 mg/1 of streptomycin sulfate) Matusick et al. 2011). Hylastes beetles and their selective nutrient agar (Jacobs & Wingfield 2001). associated fungi are thought to collectively contribEach adult insect was also rolled on CSMA to re ute to LPD by causing root damage through fungal cover ophiostomatoid fungi from insect bodies infection and insect feeding (Eckhardt et al. 2007). (Zanzot et al. 2010). Isolated ophiostomatoid fun Root and lower stem feeding beetle species (Colegal species were identified using morphological optera: Curculionidae) includingH. salebrosus and characters and established keys. Unknown iso H. tenuis are found in increased numbers within lates were sequenced and confirmed as new spe LPD-affected stands, compared to healthy stands cies by M. Wingfield (Forestry and Agricultural (Eckhardt et al. 2007, Menard 2007). A recent Biotechnology Institute, Pretoria, South Africa), study has found Hylastes to be the most common A total of 157 adult beetles (Coleoptera: Cur bark beetle genus observed in loblolly pine stands culionidae) were collected from roots, most from in central Alabama (Thompson 2011), illustrating dying trees (Table 1). A majority of the beetles col their dominance in this ecosystem. Despite their lected were Hylastes species, including a total of dominance and potential for damage in loblolly 113 H. salebrosus (all from dying trees) and 34 pine, the breeding behavior of Hylastes species in H. tenuis (31 from dying and 3 from dead trees), the southeastern United States is not well underIn addition, H. salebrosus larvae and pupae were stood. In the current study, observational methcollected from dying trees. Other adult insect spe ods were used to investigate the potential for cies collected include the black turpentine beetle

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.

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.

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.