Impact of fire on leaf nutrients,arthropod fauna and herbivory of native and exotic eucalypts in Kings Park,Perth, Western Australia

The vegetation of Kings Park, near the centre of Perth, Western Australia, once had an overstorey of Eucalyptus marginata (jarrah) or Eucalyptus gomphocephala (tuart), and many trees still remain in the bushland parts of the Park. Avenues and roadsides have been planted with eastern Australian species, including Eucalyptus cladocalyx (sugar gum) and Eucalyptus botryoides (southern mahogany), both of which have become invasive. The present study examined the effect of a recent burn on the level of herbivory on these native and exotic eucalypts. Leaf damage, shoot extension and number of new leaves were measured on tagged shoots of saplings of each tree species in unburnt and burnt areas over an 8‐month period. Leaf macronutrient levels were quantified and the number of arthropods on saplings was measured at the end of the recording period by chemical knockdown. Leaf macronutrients were mostly higher in all four species in the burnt area, and this was associated with generally higher numbers of canopy arthropods and greater levels of leaf damage. It is suggested that the pulse of soil nutrients after the fire resulted in more nutrient‐rich foliage, which in turn was more palatable to arthropods. The resulting high levels of herbivory possibly led to reduced shoot extension of E. gomphocephala, E. botryoides and, to a lesser extent, E. cladocalyx. This acts as a negative feedback mechanism that lessens the tendency for lush, post‐fire regrowth to outcompete other species of plants. There was no consistent difference in the levels of the various types of leaf damage or of arthropods on the native and the exotic eucalypts, suggesting that freedom from herbivory is not contributing to the invasiveness of the two exotic species.     

Diversity patterns of eucalypt canopy arthropods in eastern and western Australia

1. As part of a larger study on canopy arthropods and birds, a 1‐year chemical knockdown study was carried out in one Western Australian forest, where jarrah Eucalyptus marginata and marri E. (Corymbia) calophylla were sampled, and one eastern Australian (New South Wales) forest, where narrow‐leaved ironbark E. crebra and grey box E. moluccana were sampled. 2. Ten trees of each species were sampled during each of the four seasons and the arthropods were sorted to morphospecies level. This paper documents the foliage‐associated component of arboreal arthropod communities and compares arthropod species richness within orders and families, between tree species, and between the two forest types. 3. Hymenoptera, Coleoptera, Diptera, and Araneae were the richest in species. Nine hundred and seventy‐six species in 173 families were found in the eastern Australian forest, while 687 species in 176 families were found in the western Australian forest. Only 53% of families were common to both forests, but almost half the families recorded were represented by fewer than five species. Species overlap between tree species in each region was 40–53%. 4. Analysis using nonparametric bootstrapping procedures showed that sampling of foliage was comprehensive and that only 4–9% more species would be expected with more intensive sampling of the canopy. Absolute richness, as well as differences between tree species and regions, therefore appear to be real and not the result of sampling errors. As a consequence, arthropod species richness in Australian eucalypt forests is shown to be substantially greater than previous estimates.     

Jarrah Forest Restoration in Western Australia: Canopy and Topographic Effects

The restoration of the northern jarrah (Eucalyptus marginata) forest after bauxite mining is a major objective of Alcoa of Australia Limited. The typically variable and sometimes low emergence of broadcast seed of jarrah-forest plant species may relate to microclimatic changes associated with mining disturbance. This study examined the effect of the presence and absence of a canopy and topographic position in the post-mining landscape on the emergence of four canopy species (E. marginata, E. calophylla, E. patens, and E. diversicolor) and related these patterns to detailed measures of surface soil temperature and moisture. The absence of a canopy in the restoration appeared to result in adverse microclimatic conditions for the successful early establishment of E. marginata and E. calophylla from seed, particularly in the low topographic regions of the restoration. Emergence beneath a canopy compared to that in the open was 17% and 6%, respectively, for E. marginata and 23% and 2%, respectively, for E. calophylla. For both species, emergence was also greater at upland than at lowland open restoration sites (9% and 3%, respectively, for E. marginata; 4% and 0.3%, respectively, for E. calophylla). In contrast, canopy removal and position on the topographic landscape did not reduce the early establishment success of E. patens and E. diversicolor. Field measurements revealed that soils were drier and that diurnal temperature fluctuations were wider in the open restoration sites than beneath a canopy. Furthermore, cold conditions were more frequent at lowland than at upland restoration sites, suggesting the occurrence of cold-air drainage to Jew-lying areas. It is therefore possible that the field emergence patterns reflected the lower tolerance of E. marginata and E. calophylla than both E. diversicolor and E. patens to cold and dry surface-soil conditions. The ecological significance and practical implications of the results are discussed.     

Seasonality of canopy invertebrate communities in eucalypt forests of eastern and western Australia

Abstract Chemical knockdown procedures were used to sample canopy arthropods at 3 month intervals over 1 year at two sites, one in eastern Australia and the other in western Australia. Samples were taken from narrow-leaved ironbark, Eucalyptus crebra, and grey box, Eucalyptus moluccana, in the east and from jarrah, Eucalyptus marginata, and marri, Eucalyptus calophylla, in the west. Arthropods were more abundant on trees in eastern Australia and exhibited different seasonal patterns from those in the west. Members of different functional groups exhibited different seasonal patterns, with some herbivorous groups responding to times of leaf production, decomposers and fungus feeders responding to high moisture availability, and predators/parasites responding to the abundance of food items. Seasonal variability was slightly higher in the west, possibly reflecting the greater seasonal amplitude in rainfall. In the eastern forest, proportionately more taxa peaked in spring or summer and declined to minimum numbers in winter. In the western forest several taxa attained peak numbers in autumn, winter or spring, while others declined to minimum values in winter or summer. The phenological patterns of canopy arthropods appear to be linked to the condition of the host plant and/or to climatic factors. Comparison of the western Australian data to those from a second year of sampling at a time when rainfall was greater and fell later into the season indicated that variability in arthropod numbers between years can be as great as that between seasons. Implications of the variability in seasonal and annual patterns of canopy invertebrate communities are discussed in relation to the need for long-term sampling and in relation to evaluating the impact of disturbance on forest communities.     

Differences in nitrogen use efficiency between leaves from canopy and subcanopy trees

We examined the effects of increasing light availability along a vertical gradient within a forest community on the efficiency of leaf nitrogen (N) use in individual trees. The N contents of green and senescent leaves in canopy and subcanopy trees of an evergreen coniferous species, Podocarpus nagi, and an evergreen hardwood species, Neolitsea aciculata, were analyzed in a mixed forest community at Mt Mikasa, Nara City, Japan. The inverse of N concentration (N_C) in senescent leaves was used as an index of N use efficiency (NUE) at the leaf-level. The leaf-level NUE was higher in canopy trees than in subcanopy trees in both P.nagi and N.aciculata, although soil N mineralization rates around canopy and subcanopy trees did not differ significantly. The N_C in green leaves was lower in canopy trees than in subcanopy trees. The ratio of resorbed N in senescent leaves to the N content in green leaves was higher in canopy trees than in subcanopy trees. The higher leaf-level NUE of canopy trees was partly a result of lower N_C in living tissues and partly because of greater N resorption during senescence. The present study suggested that the leaf-level NUE could be increased in response to an imbalance between soil N and light availability caused by spatial community structure.     

The impact of defoliation on the foliar chemistry of southern rātā (

Brushtail possums (Trichosurus vulpecula) tend to eat young canopy foliage in southern rätä (Metrosideros umbellata), and browsing tends to be concentrated on only a few trees. Samples collected as part of an artificial defoliation experiment were analysed for NPK (nitrogen, phosphorus, and potassium), carbohydrate, and polyphenolic concentrations to determine whether changes in foliar chemistry associated with defoliation provide an explanation for these patterns of browsing. In non-defoliated trees, NPK concentrations were highest in young leaves and declined with age, while concentrations of carbohydrates and polyphenolics were independent of leaf age. Nitrogen, phosphorus and polyphenolic concentrations were consistently higher in canopy (sun) versus subcanopy (shade) foliage regardless of leaf age, a trend that was reversed for potassium. Partial (50%) defoliation had little effect on foliar chemistry, regardless of its timing. Total (100%) defoliation stimulated NPK concentrations and depressed condensed tannin concentrations of new foliage produced by the surviving shoots. These results suggest that brushtail possums may focus their feeding on only a few trees because of nutritional changes to leaves as a result of browsing.     

Invertebrate communities on Western Australian eucalypts: A comparison of branch clipping and chemical knockdown procedures

Chemical knockdown and branch clipping procedures were used in wandoo (Eucalyptus wandoo) woodland and jarrah (E. marginata)/marri (E. calophylla) open-forest to sample arboreal invertebrate faunas on three species of Western Australian eucalypts. Jarrah was sampled in both habitats and had significantly lower invertebrate populations and a less diverse fauna than either wandoo or marri. The two procedures provided similar results with respect to the relative abundance of invertebrates on each plant species but the knockdowns sampled a more diverse fauna, including species sheltering in or on bark. Chemical knockdowns underestimated the abundance of sessile invertebrates, such as psyllids. Branch clipping sampled insufficient numbers of large, mobile, or cryptic invertebrates to estimate abundances, but provided a more accurate estimate of the abundance of sessile, leaf-dwelling organisms. Neither procedure provides a complete sample of arboreal invertebrates, but they are complementary. When used in conjunction with each other a more complete estimate of arboreal invertebrate abundance and diversity is obtained. Both procedures can be used concurrently with only a small increase in field time.     

Responses of forest eucalypts to moderate and high intensity fire in the Tingle Mosaic,south‐western Australia: comparisons between locally endemic and regionally distributed species

Responses of three locally endemic (Eucalyptus brevistylis, Eucalyptus jacksonii and Eucalyptus guilfoylei) and three co‐occurring regional eucalypts (Eucalyptus marginata, Eucalyptus diversicolor and Corymbia calophylla) to moderate‐ and high‐intensity fires were examined in granitic terrain of the Tingle Mosaic, south‐western Australia. Significant associations between diameter distributions and community type (CT) for each species (P < 0.001) suggest that fire response will also vary according to the habitat/fire interaction. None of the species were fire sensitive, although responses differed both within and between species, and with CT. All species examined predominately consisted of several cohorts of regeneration within a forest stand. Each species had thick bark and re‐sprouted from crown epicormics following 100% scorch of the mature tree. The quantity and type of regeneration in relation to gaps created by individual dead trees following fire differed between species; for example, E. guilfoylei regeneration was strongly associated with gaps, and C. calophylla with non‐gaps. However, regeneration of the two tall open‐forest species, E. jacksonii and E. diversicolor were not most associated with either gaps or non‐gaps. The very low levels of regeneration of E. brevistylis following fire and the high proportion of stems of E. jacksonii that were hollow butted (40% of stems > 1 m DBHOB) may be factors associated with narrow endemism of these species and may affect the vulnerability of these eucalypts to fire. The interaction of seed availability, intense fires and subsequent rainfall may be critical in the long term survival of these species. Eucalyptus guilfoylei, by contrast, appears well adapted to the increasing levels of disturbance likely in the region where these species occur. The vulnerability of a locally endemic species in a fire‐prone environment is likely to reflect differences to the prevailing adaptations of the dominant species rather than an inherent ability of the species to survive or respond. Management regimes must account for variations in species responses to fire in different CTs if the long‐term survival chances of local endemics are to be enhanced.     

Nitrogen fertilization enhances water‐use efficiency in a saline environment

Effects of salinity and nutrients on carbon gain in relation to water use were studied in the grey mangrove, Avicennia marina, growing along a natural salinity gradient in south‐eastern Australia. Tall trees characterized areas of seawater salinities (fringe zone) and stunted trees dominated landward hypersaline areas (scrub zone). Trees were fertilized with nitrogen (+N) or phosphorus (+P) or unfertilized. There was no significant effect of +P on shoot growth, whereas +N enhanced canopy development, particularly in scrub trees. Scrub trees maintained greater CO₂ assimilation per unit water transpired (water‐use efficiency, WUE) and had lower nitrogen‐use efficiency (NUE; CO₂ assimilation rate per unit leaf nitrogen) than fringe trees. The CO₂ assimilation rates of +N trees were similar to those in other treatments, but were achieved at lower transpiration rates, stomatal conductance and intercellular CO₂ concentrations. Maintaining comparable assimilation rates at lower stomatal conductance requires greater ribulose 1·5‐bisphosphate carboxylase/oxygenase activity, consistent with greater N content per unit leaf area in +N trees. Hence, +N enhanced WUE at the expense of NUE. Instantaneous WUE estimates were supported by less negative foliar δ¹³C values for +N trees and scrub control trees. Thus, nutrient enrichment may alter the structure and function of mangrove forests along salinity gradients.     

Ecophysiology of Eucalyptus marginata and Corymbia calophylla in decline in an urban parkland

Eucalypt trees are in decline throughout urban landscapes of south western Australia. This study investigated the cause of decline in Eucalyptus marginata and Corymbia calophylla trees in parkland and compared water and nutrient relations with healthy trees in adjacent bushland in Perth, Western Australia. It was hypothesized that: (i) trees were drought stressed through competition for soil water by the vigorous turf; (ii) excessive uptake of nitrogen, because of fertilizer application to turf, caused toxicity; and/or (iii) micronutrient (Cu, Fe, Mn and/or Zn) deficit was induced by high‐pH irrigation water applied to turf around parkland trees. Leaf water potential showed aseasonal variation in the irrigated parkland trees and foliar δ¹³C indicated that parkland trees generally had low water‐use efficiency and were not drought stressed relative to bushland trees. Foliar N levels were not significantly different between parkland and bushland trees indicating that excess N uptake was not a factor in the decline. Foliar total Fe, ‘metabolically active’ Fe, Cu and Zn concentrations were not significantly different between parkland and bushland trees. Foliar manganese concentrations were indicative of deficiency and significantly lower in parkland trees (5–14 µg g^?1) relative to bushland trees (22–35 µg g^?1). It is concluded that application of alkaline irrigation water to the parkland site reduced the plant‐availability of Mn; however, our study of only one parkland site does not allow us to generalize the results across other parklands.     

Foliar and twig macronutrients (N,P, K,Ca and Mg) in selected species ofEucalyptus used in rehabilitation: sources of variation

Summary The macronutrient variation within four 6 year oldEucalyptus saligna and four 5 year oldE. wandoo growing on rehabilitated bauxite pits was determined. Significant differences in mean nutrient concentrations were generally recorded between good soil condition sites and poor soil sites, between tree individuals, branch height, and plant organ type; but mean nutrient values were not different among canopy aspects. Fully expanded leaves of the current year provided the most uniform nutrient levels among the plant organs and showed major differences between sites with good soil nutrient conditions and those with poor conditions. Differences in foliar and branch levels of N, P, K, Ca and Mg, the variation between sites, canopy heights and plant organ types, and the use of foliar nutrient levels to indicate deficiencies are discussed.     

Gene flow of the canker pathogen Botryosphaeria australis between Eucalyptus globulus plantations and native eucalypt forests in Western Australia

Abstract The eucalypt plantation industry in Western Australia provides a unique opportunity to study the movement of pathogens between closely related host taxa. Eucalyptus globulus, a native to Tasmania and south‐eastern Australia, is the predominant species in Western Australian plantations, often being planted adjacent to native forest containing Eucalyptus marginata and Eucalyptus diversicolor. Since the commencement of the plantation industry 20 years ago, several fungal species, previously known only to eastern Australia or overseas, have been reported on E. globulus in Western Australia. Botryosphaeria australis is a newly described species, recently found causing cankers on Acacia spp. in eastern Australia. However, during a routine survey, B. australis was found to be the predominant species associated with E. globulus plantations and native Eucalyptus spp. in Western Australia. In this study, six short simple repeat markers were used to evaluate genetic diversity and gene flow between collections of B. australis from native eucalypt forest and E. globulus plantations at two locations in south‐western Australia. In both cases, there was no restriction to gene flow between the plantations and the adjacent native forest. Botryosphaeria australis has now been isolated from a wide range of hosts across south‐western Australia and was not isolated from E. globulus in Tasmania or South Australia. This extensive distribution and host range suggests B. australis is native to Western Australia. This study demonstrates the ability of a pathogen to move between plantation and forests.     

Changes in structure of over‐ and midstory tree species in a Mediterranean‐type forest after an extreme drought‐associated heatwave

Worldwide, extreme climatic events such as drought and heatwaves are associated with forest mortality. However, the precise drivers of tree mortality at individual and stand levels vary considerably, with substantial gaps in knowledge across studies in biomes and continents. In 2010–2011, a drought‐associated heatwave occurred in south‐western Australia and drove sudden and rapid forest canopy collapse. Working in the Northern Jarrah (Eucalyptus marginata) Forest, we quantified the response of key overstory (E. marginata, Corymbia calophylla) and midstory (Banksia grandis, Allocasuarina fraseriana) tree species to the extreme climate event. Using transects spanning a gradient of drought impacts (minimal (50–100 m), transitional (100–150 m) and severe (30–60 m)), tree species mortality in relation to stand characteristics (stand basal area and stem density) and edaphic factors (soil depth) was determined. We show differential mortality between the two overstory species and the two midstory species corresponding to the drought‐associated heatwave. The dominant overstory species, E. marginata, had significantly higher mortality (~19%) than C. calophylla (~7%) in the severe zone. The midstory species, B. grandis, demonstrated substantially higher mortality (~59%) than A. fraseriana (~4%) in the transitional zone. Banksia grandis exhibited a substantial shift in structure in response to the drought‐associated heatwave in relation to tree size, basal area and soil depth. This study illustrates the role of climate extremes in driving ecosystem change and highlights the critical need to identify and quantify the resulting impact to help predict future forest die‐off events and to underpin forest management and conservation.     

Seasonal changes in foliar macronutrients (N,P, K,Ca and Mg) inEucalyptus saligna Sm. andE. wandoo Blakely growing in rehabilitated bauxite mine soils of the Darling Range,Western Australia

Summary Seasonal changes in the foliar concentration of macronutrients (N, P, K, Ca and Mg) in sapling trees ofEucalyptus saligna Sm. andE. wandoo Blakely growing in rehabilitated bauxite mined areas in the Darling Range of Western Australia are described. Foliar N concentration decreased with age of the fully expanded leaf tissue. Leaf N concentrations were also high when rates of litter decomposition were expected to be high during the period of early spring. The greatest foliar N difference between trees growing in good soil conditions and those from poorer soil conditions also occurred during this period. Levels of P in leaves were highest in young developing leaves but once the leaves reached full size, no seasonal trend in P concentration was observed. Foliar K was lower during the winter and probably related to the period of maximum leaching by precipitation. High foliar K during summer, however, could be related to the role of K in lowering cellular water potential. Leaf Ca was highest during early sping. Low mobility of cellular Ca during the cool portion of the year was indicated. Foliar Mg showed a weak pattern of decreasing concentration with leaf age. The best season for sampling for these broadleafed evergreen species to provide information on plant nutrient status appears to be in spring.     

Comparison of the nutrient ecology of coastal Banksia grandis elfinwood (windswept shrub‐like form) and low trees,Cape Leeuwin‐Naturaliste National Park,Western Australia

Abstract Trees growing along windy coasts often have canopies that are greatly reduced in size by the sculpting effects of wind and salt spray. Trees with environmentally reduced stature are called elfinwood (windswept shrub‐form or krummholz) and are ecologically important because they represent outposts growing at the limit of tree success. The purpose of this study was to assess if Banksia grandis elfinwood growing at Cape Leeuwin had a different nutrient status than normal low‐form (LF) trees growing nearby, and if nutrient deficiencies, toxicities and/or imbalances were among the limiting factors imposed on elfinwood. The concentrations of N, P, K, Ca, Mg, Na, Cl^–, Fe, Mn, Zn, Cu, Mo and B were analysed for mature green foliage, immature foliage, foliage litter, flowers and soil. When the elfinwood and LF trees were compared, the foliar nutrient status was generally similar, except that elfinwood foliage had significantly higher mean concentrations of N, Zn and Cu, while LF trees had higher Fe and Mn contents. Many nutrients were conserved before leaves were shed in both elfinwood and LF trees, including N, P, K, Na, Cl^–, Mn and Cu (LF trees also conserved Ca and Mg). However, elfinwood and LF tree‐litter contained significantly higher Fe concentrations than green foliage (elfinwood litter also had higher levels of Mg and B). It is tempting to suggest that the translocation of Fe into leaves before they were shed is a regulation mechanism to prevent Fe toxicity, or imbalance in the Fe : Mn ratio. Proteoid roots strongly acidify the soil to mobilize P, which also chemically reduces Fe⁺³ to plant‐available Fe⁺². The increased supply of Fe⁺² in the rhizosphere, caused by the action of proteoid roots, might tend to defeat self‐regulation of Fe uptake. It is possible that excess Fe accumulation in the plant might be regulated, in part, by exporting Fe into the leaves before they are shed. The nutrient status of B. grandis elfinwood is compared with mountain elfinwood of North America. The extreme habitat of coastal elfinwood provides many theoretical pathways for nutrient limitation, but B. grandis elfinwood at Cape Leeuwin does not appear to be nutrient deficient.     

Changes in Mineral Content of Eucalyptus marginata and E. calophylla Grown under Controlled Conditions and Inoculated with Phytophthora cinnamomi

Mineral concentrations in infected roots and shoots were compared with similar material, but pathogen free, for the susceptible Eucalyptus marginata and the field resistant E. calophylla. All plants were mycorrhiza-free, grown under controlled conditions and inoculated with an axenic suspension of zoospores of Phytophthora cinnamomi. Plant material was ashed 30 days after inoculation and analyzed in an external proton beam using PIXE and nuclear reaction analyis to detect differences in mineral concentrations. The mineral content of infected roots of E. marginata was below that of the uninfected roots for all elements detected except chlorine, chromium and rubidium. The reductions were significant for calcium and copper. No such reduction was found, in E. calophylla roots, but some changes were detected. Shoot: root ratios of E. marginata showed significant differences associated with infection in phosphorus, calcium, copper and zinc. Relatively large differences were also recorded for sulphur chlorine and potassium. Shoot: root ratios of infected E. calophylla showed fewer differences but contained only half the concentrations of sulphur and potassium present in the controls. The reduced mineral concentrations reflect the failure in conduction of water and minerals through the infected and susceptible root system.     

Sheoak woodlands: a newly identified habitat for western ringtail possums

Although critically endangered western ringtail possums (Pseudocheirus occidentalis) have been recorded from a variety of vegetation types in southwestern Australia, the extent to which many of these vegetation types are occupied by the species remains unknown. We conducted spotlight surveys for western ringtail possums between March and April 2018 in Albany, southwestern Australia, to determine the species’ occupancy in 2 vegetation types. Using occupancy models, we demonstrated that sheoak (Allocasuarina fraseriana) woodlands, previously unrecognized as providing western ringtail possum habitat, support the species, although the median probability of their occupancy was lower than in marri (Corymbia calophylla) and eucalypt (Eucalyptus marginata and E. staeri) woodlands. Use of trees and other habitat components by western ringtail possums varied within and between vegetation types. Sheoak woodlands are likely critical for western ringtail possums and given the apparent flexibility in vegetation types used by the species, investigation of the potential value of other vegetation types for this species has conservation merit. © 2019 The Wildlife Society.     

Phytophagous insects enhance nitrogen flux in a desert creosotebush community

Summary We tested the hypothesis that herbivorous insects on desert shrubs contribute to short-term nitrogen cycling, and increase rates of nitrogen flux from nutrient rich plants. Creosotebush (Larrea tridentata) shrubs were treated with different combinations of fertilizer and water augmentations, resulting in different levels of foliage production and foliar nitrogen contents. Foliage arthropod populations, and nitrogen in canopy dry throughfall, wet throughfall and stemflow were measured to assess nitrogen flux rates relative to arthropod abundances on manipulated and unmanipulated shrubs over a one-month period during peak productivity. Numbers and biomass of foliage arthropods were significantly higher on fertilized shrubs. Sap-sucking phytophagous insects accounted for the greatest numbers of foliage arthropods, but leaf-chewing phytophagous insects represented the greatest biomass of foliage arthropods. Measured amounts of bulk frass (from leaf-chewing insects) were not significantly different among the various treatments. Amounts of nitrogen from dry and wet throughfall and stemflow were significantly greater under fertilized shrubs due to fine frass input from sap-sucking insects. Increased numbers and biomass of phytophagous insects on fertilized shrubs increased canopy to soil nitrogen flux due to increased levels of herbivory and excrement. Nitrogen excreted by foliage arthropods accounted for about 20% of the total one month canopy to soil nitrogen flux, while leaf litter accounted for about 80%.     

Hemlock Woolly Adelgid in New England Forests: Canopy Impacts Transforming Ecosystem Processes and Landscapes

Exotic insect pests may strongly disrupt forest ecosystems and trigger major shifts in nutrient cycling, structure, and composition. We examined the relationship between these diverse effects for the hemlock woolly adelgid (HWA, Adelges tsugae Annand) in New England forests by studying its impacts on local canopy processes in stands differing in infestation levels and linking these impacts to shifts in canopy nutrient cycling and stand and landscape effects. HWA initiated major changes in canopy biomass and distribution. Whereas uninfested trees exhibit a significant decline in canopy biomass from the center to the periphery and a positive correlation between total needle litter and estimated biomass, infested trees have significantly less total canopy biomass, produce less new foliage, shed relatively more needles, and exhibit no correlation between litter and canopy biomass. Foliar N content of infested trees was 20%–40% higher than reference trees, with the strongest increase in young foliage supporting the highest densities of HWA. Foliar %C was unaffected by HWA or foliar age. Epiphytic microorganisms on hemlock needles exhibited little variation in abundance within canopies, but colony-forming units of bacteria, yeast, and filamentous fungi were 2–3 orders of magnitude more abundant on medium and heavily infested than uninfested trees. Throughfall chemistry, quantity, and spatial pattern were strongly altered by HWA. Throughfall exhibits a strong gradient beneath uninfested trees, decreasing in volumes from the canopy periphery to the trunk by more than 45%. The amount of throughfall beneath infested trees exhibits no spatial pattern, reaches 80%–90% of the bulk precipitation, and is characterized by significantly higher concentrations of nitrogen compounds, dissolved organic carbon, and cations. Across the southern New England landscape there is a strong south-to-north gradient of decreasing hemlock tree and sapling mortality and understory compositional change that corresponds to the duration of infestation. Regionally, black birch (Betula lenta L.) is profiting most from hemlock decline by significantly increasing in density and cover. These findings suggest that it is necessary to study the connections between fast/small-scale processes such as changes in nutrient cycling in tree canopies and slow/integrative processes like shifts in biogeochemieal cycling and compositional changes at forest stands and landscapes to better understand the effects of an exotic pest species like HWA on forest ecosystem structure and function.     

Calcium addition at the Hubbard Brook Experimental Forest increases the capacity for stress tolerance and carbon capture in red spruce (Picea rubens) trees during the cold season

Red spruce (Picea rubens Sarg.) trees are uniquely vulnerable to foliar freezing injury during the cold season (fall and winter), but are also capable of photosynthetic activity if temperatures moderate. To evaluate the influence of calcium (Ca) addition on the physiology of red spruce during the cold season, we measured concentrations of foliar polyamines and free amino acids (putative stress-protection compounds), chlorophyll (a key photosystem component), and sapwood area (a proxy for foliar biomass), for trees in Ca-addition (CaSiO₃ added) and Ca-depleted (reference) watersheds at the Hubbard Brook Experimental Forest (NH, USA). Ca-addition increased concentrations of the amino acids alanine and γ-aminobutyric acid (GABA) and the polyamines putrescine (Put) and spermidine (Spd) in November, and Put in February relative to foliage from the reference watershed. Consistent with increased stress protection, foliage from the Ca-addition watershed had higher total chlorophyll and chlorophyll a concentrations in February than foliage from the reference watershed. In contrast, foliage from the reference watershed had significantly lower glutamic acid (Glu) and higher alanine (Ala) concentrations in February than foliage from the Ca-addition watershed. Imbalances in Ala:Glu have been attributed to cold sensitivity or damage in other species. In addition to concentration-based differences in foliar compounds, trees from the Ca-addition watershed had higher estimated levels of foliar biomass than trees from the reference watershed. Our findings suggest that Ca-addition increased the stress tolerance and productive capacity of red spruce foliage during the cold season, and resulted in greater crown mass compared to trees growing on untreated soils.