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Past work shows a significant negative correlation between foliar oregonin concentration and western tent caterpillar (Malacosoma californicum Packard) feeding on red alder (Alnus rubra Bong.). Above an oregonin threshold of 20% leaf dry weight, little feeding by caterpillars is observed. Concentrations of defensive chemicals are influenced by plant genotype, environmental conditions, insect feeding, and the interactions of these factors. Our objective was to measure the effects of nitrogen (N) availability and wounding on foliar oregonin and condensed tannin concentrations in red alder genotypes. One-year-old seedlings from 100 half-sib red alder families were treated with two levels of ammonium nitrate (NH4NO3) for two growing seasons in a common garden. In the second year, leaves from 50 families from the fertilization experiment were used in a bioassay feeding experiment to determine the effects of N fertilization and genotype on WTC damage, and to identify a subset of 20 families with a range of damage to analyze for phytochemical composition. In separate experiments, wound-induction treatments were conducted outdoors and, in a greenhouse using the N treated trees in their third and fourth year, respectively. Foliar condensed tannin, oregonin and N concentrations were measured and ranked among the plant genotypes, and between the two N treatments and two wounding treatments. Results showed that oregonin and condensed tannin concentrations varied among the alder genotypes. Leaf N concentration was negatively correlated with concentration of oregonin. Neither of the measured phenolic compounds responded to wounding. The results suggest that red alder foliar oregonin and condensed tannin are likely constitutive defenses that are largely determined by genotype, and that the negative correlation of defense compounds with plant internal N status holds in this N-fixing tree.

Red alder (Alnus rubra Bong.) is a tree of high economic and ecological importance, yet is subject to severe defoliation during episodic outbreaks of tent caterpillars (Malacosoma spp.). We evaluated variation in western tent caterpillar (Malacosoma californicum Packard, 1864) (WTC) resistance among and within red alder populations and clones and investigated potential defense mechanisms. Bioassay feeding trials were conducted with WTC on 20 red alder clones from 10 provenances (two clones per provenance). Phenology and quality of red alder leaves were analyzed to determine whether bud burst, leaf chemical content, water content, or physical traits are determinants of WTC preference. Alder clones differed in percent leaf area eaten by WTC and in leaf defense traits. The concentrations of total phenolics, condensed tannin, and the diarylheptanoid oregonin were negatively correlated with the percent leaf area eaten by caterpillars. In addition, a potential threshold was observed above which the concentration of each of the chemicals appeared to reduce WTC feeding. In particular, foliar concentrations of oregonin >20% of leaf dry mass were consistently associated with reduced feeding. The effects of oregonin concentration in red alder leaves on tent caterpillar feeding is a novel finding.

Small inreases in CO2 stimulate nitrogen fixation and plant growth. Increasing soil N can inhibit nitrogen fixation. However, no studies to date have tested how nitrogen fixing plants perform under ancient CO2 levels (100 MYA), when nitrogen fixing plants evolved, with different levels of N additions. The aim of this study was to assess if ancient CO2, compared to present, favors nitrogen fixers over a range of soil nitrogen concentrations. Nitrogen fixers (Alnus incana ssp. rugosa, Alnus viridis ssp. crispa, and Alnus rubra) and their close non-nitrogen fixing relatives (Betula pumila, Betula papyrifera, Betula glandulosa) were grown at ancient (1600 ppm) or present (400 ppm) CO2 over a range of soil N levels, equivalent to 0, 10, 50, and 200 kg N ha−1 year−1. The growth of non-N fixing plants increased more than N fixing plants in response to the increasing N levels. When grown at an ancient CO2 level, the N level at which non-nitrogen fixing plant biomass exceeded nitrogen fixing plant biomass was twice as high (61 kg N ha−1 year−1) as the N level when plants were grown at the ambient CO2 level. Specific nodule activity was also reduced with an increasing level of soil N. Our results show there was a greater advantage in being a nitrogen fixer under ancient levels of CO2 compared with the present CO2 level.

Plants synthesize a wide range of bioactive secondary metabolites to defend against pests and pathogens. Red alder (Alnus rubra) bark, root, and leaf extract have a long history of use in traditional medicine and hygiene. Diarylheptanoids, especially oregonin ((5S)-1,7-bis(3,4-dihydroxyphenyl)-5-(β-D-xylopyranosyloxy)-heptan-3-one), have been identified as major bioactive constituents. Diarylheptanoids have become a focus of research following reports of their antioxidant, antifungal, and anti-cancer activities. Recent data suggest that high oregonin concentration is associated with resistance of red alder leaves to western tent caterpillar (Malacosoma californicum) defoliation. Here we test effects of this compound directly on leaf-eating insects. Purified oregonin was examined in insect choice and toxicity tests using lepidopteran caterpillars. The compound exhibited significant anti-feedant activity against cabbage looper (Trichoplusia ni), white-marked tussock moth (Orgyia leucostigma), fall webworm (Hyphantria cunea), and M. californicum at concentrations corresponding to oregonin content of the most resistant alder clones in previous experiments. Toxicity tests were carried out with cabbage looper larvae only, but no contact or ingested toxicity was detected. Our results suggest that oregonin at levels found in red alder leaves early in the growing season may contribute to protecting red alder from leaf-eating insects.

Introduction The diarylheptanoid xyloside oregonin ((5S)-1,7-bis(3,4-dihydroxyphenyl)-5-(β-d-xylopyranosyloxy)-heptan-3-one) has significant medicinal potential and is found at high concentration in leaves and bark of red alder (Alnus rubra). Objectives To establish inexpensive and easily scaled methods for the extraction and purification of oregonin from timber by-products. Methods We developed a method combining aqueous extraction with spray drying of red alder extract into a powder, thus reducing the need for organic solvents used in traditional Soxhlet extraction or in solvent partitioning. Flash chromatography was utilised to purify oregonin from crude spray-dried alder extract. Results Crude spray-dried alder extract was comprised of an average of 9% of the diarylheptanoid compound oregonin. Less than 10% thermal degradation of oregonin was observed using extraction temperatures between 25°C and 50°C, followed by spray drying. The structure of purified oregonin was validated using high-performance liquid chromatography (HPLC), mass spectrometry (MS), ultraviolet spectroscopy (UV), and nuclear magnetic resonance (NMR). Conclusion The developed method was robust, repeatable, and yielded purified oregonin of greater than > 95% purity (average of 95.8%). Our analysis represents the most complete NMR characterisation of oregonin reported to date.

Red alder (Alnus rubra [Bong.]), is the most common broadleaf species in the Pacific Northwest, and is widely distributed as a component of mixed stands with conifers. It rapidly overtops associated juvenile conifers and is a strong competitor. However, red alder is also a symbiotic nitrogen fixer and can contribute to improvement in soil nitrogen, as well as serving other roles such as improving biodiversity and forest health. We examined data collected in mixed red alder-conifer plantations established between 1992 and 1999 to examine the effects of red alder on the growth of Douglas-fir (Pseudotsuga menziessi (Mirb.) Franco, Fd) and western redcedar (Thuja plicata Donn., Cw). Results suggest that densities of up to 400 alder trees per hectare may be acceptable and possibly desirable on nitrogen deficient sites. The growth of Fd may be enhanced when red alder density is relatively low at older ages. We found that the competitive effects of red alder on conifer growth were consistently lower than that of the conifers (Fd and Cw) while alder had stronger effects on light availability. The relationship between volume growth and competition indices varied between the two conifer species: (1) for Fd the highest correlations were obtained using a distance-dependent competition index based on the height differences, and (2) for Cw the highest correlations were with distance-independent Lorimer’s index. As also expected due to Cw’s greater shade tolerance, its volume growth showed less sensitivity to the presence of red alder than Fd.

Practical methods for tree species identification are important for both land management and scientific inquiry. LiDAR has been widely used for species mapping due to its ability to characterize 3D structure, but in structurally complex Pacific Northwest forests, additional research is needed. To address this need and to determine the feasibility of species modeling in such forests, we compared six approaches using five algorithms available in R’s lidR package and Trimble’s eCognition software to determine which approach most consistently identified individual trees across a heterogenous riparian landscape. We then classified segments into Douglas fir (Pseudotsuga menziesii), black cottonwood (Populus balsamifera ssp. trichocarpa), and red alder (Alnus rubra). Classification accuracies based on the best-performing segmentation method were 91%, 92%, and 84%, respectively. To our knowledge, this is the first study to investigate tree species modeling from LiDAR in a natural Pacific Northwest forest, and the first to model Pacific Northwest species at the landscape scale. Our results suggest that LiDAR alone may provide enough information on tree species to be useful to land managers in limited applications, even under structurally challenging conditions. With slight changes to the modeling approach, even higher accuracies may be possible.

Populus trichocarpa Torr. and Gray (black cottonwood) is an economically and ecologically important tree species native to western North America. It serves as a model tree species in biology and genetics due to its relatively small genome size, rapid growth, and early reproductive maturity (Jansson and Douglas 2007). Black cottonwood is susceptible to root rot caused by at least one species of Armillaria (Raabe 1962), a globally distributed genus that exhibits diverse ecological behaviors (Klopfenstein et al. 2017) and infects numerous woody plant species (Raabe 1962). However, several Armillaria spp. have been isolated from Populus spp. in North America (Mallet 1990), and the most recent report of Armillaria on P. trichocarpa used the now ambiguated name A. mellea (Vahl.) Quel. (see Raabe 1962). In April 2016, mycelial fans and rhizomorphs of an unknown Armillaria species (isolate WV-ARR-3) were collected from P. trichocarpa in a riparian hardwood stand ca. 5.5 km east of Springfield, Oregon, USA (44°3'21.133"N, 122°49'39.935"W). The host was dominant in the canopy, large in diameter (ca. 90-cm dbh) relative to neighboring trees, and exhibited minimal crown dieback (ca. < 5%). A mycelial fan was observed destroying living cambium beneath the inner bark, indicating pathogenicity. The isolate was cultured on malt extract medium (3% malt extract, 3% dextrose, 1% peptone, and 1.5 % agar) and identified as A.cepistipes on the basis of somatic pairing tests and translation elongation factor 1α (tef1) sequences (GenBank Accession No. MK172784). DNA extraction, PCR, and tef1 sequencing followed protocols of Elias-Roman et al. (2018). From nine replications of somatic incompatibility tests (18 tester isolates representing six North American Armillaria spp.), the isolate showed high intraspecific compatibility (colorless antagonism) with three A. cepistipes tester isolates (78%), but low compatibility with the other Armillaria spp. (0 - 33%) that occur in the region. Isolate WV-ARR-3 yielded tef1 sequences with a 99% identity to A. cepistipes (GenBank Accession Nos. JF313115 and JF313121). A second isolate (WV-ARR-1; GenBank Accession No. MK172783) with a nearly identical sequence was collected from a maturing P. trichocarpa in a riparian stand ca. 8 km northeast of Monroe, Oregon (44°21'47.57"N, 123°13'14.415"W) along the Willamette River, downstream from the McKenzie river tributary where WV-ARR-3 was collected. Armillaria cepistipes has been reported on Alnus rubra (red alder) in Washington, USA (Banik et al. 1996) and on broad-leaved trees in British Columbia, Canada (Allen et al. 1996). It is generally considered to be a weak pathogen on broad-leaved trees in the Pacific Northwest, but it is also associated with pathogenicity on both coniferous and deciduous trees in Europe (e.g., Lygis et al. 2005). However, a recent phylogenetic study suggested that North American A. cepistipes is phylogenetically distinct from Eurasian A. cepistipes (Klopfenstein et al. 2017), butadditional studies are needed to determine the formal taxonomic status of North American A. cepistipes. To our knowledge, A. cepistipes has not been previously confirmed on P. trichocarpa in the U.S.A. or formally reported as a pathogen of any Populus species in North America. Continued studies are needed to determine the distribution, host range, and ecological role of A. cepistipes in riparian forests of the Pacific Northwest, while monitoring its populations under changing climates.

Two sources of nitrogen to Pacific Northwest riparian areas are marine-derived nitrogen (MDN) via anadromous Pacific salmon (Oncorhynchus spp.) and atmospheric nitrogen fixation via red alder (Alnus rubra). The recent removal of two large dams on the Elwha River, WA, opened up about 60 km of previously inaccessible river habitat for anadromous salmon. We used naturally abundant stable nitrogen isotopes to establish baseline data to monitor the influx of MDN to riparian zones of Elwha River tributaries post dam removal. We sampled riparian soil, overstory, and understory vegetation in sites with nitrogen-fixing A. rubra and sites with bigleaf maple (Acer macrophyllum) at an undammed reference site, and along three tributaries, one between the former dams (accessible to anadromous salmon since 2012) and the others upstream of the former dams (no anadromous salmon). Based on δ15N measurements of soil and vegetation, we did not detect MDN at any of the tributaries, including the reference tributary. However, the understory riparian vegetation between the former dams had a higher δ15N than the other tributaries, which may be due to upstream anthropogenic nitrogen sources. Although A. rubra foliage was isotopically distinct from A. macrophyllum, and A. rubra litter had higher total nitrogen, soil and understory vegetation in A. rubra and A. macrophyllum sites did not differ isotopically. Monitoring of these areas and those further upstream on the Elwha River will allow us to trace the return of MDN to the watershed, and help to clarify the role that anadromous fish play in riparian ecosystems.

Exploratory hybridization of red alder (Alnus rubra Bong.) and white alder (Alnus rhombifolia Nutt.) was conducted to initiate a long-term investigation into the inter-specific taxon’s potential for biomass production. Red alder selections originated in stands west of Oregon’s Cascade Mountains, while white alder came from provenances west and east of the Cascades. White alder’s heretofore unknown chromosome number was determined equal to red alder (2n = 2x = 28) and included a pair of satellite chromosomes. An experimental crossing technique was proven, establishing the species’ reproductive affinity for the first time. Hybridization was substantially more productive when red alder served as the female parent as opposed to white alder. Hybridity was confirmed using four species-specific SSR markers developed from transcriptome resources for identifying sib relationships in domesticated populations. To begin a description of the inter-specific taxon, hybrids and their open-pollinated siblings were assessed at ages four through six for morphological and phenological traits and carbon and nitrogen isotopic ratios in a field planting in western Washington, USA. Principal component analysis of quantitative leaf and internode morphologies showed that red alder and hybrid alder are relatively comparable and both are distinct from white alder. Qualitative differences in leaf morphology are more diagnostic with inter-specific hybrids often intermediate to red and white alder. Principal coordinate analysis of molecular data cleanly separated the three taxa. Plans for continued investigation of the taxon as a biomass crop species are discussed including a potential role for inter-specific hybrids in adapting red alder to the effects of climate change.

In the Pacific Northwest, Alnus rubra Bong. (red alder) is a common deciduous tree species especially prevalent in riparian corridors and disturbed sites, including metropolitan areas undergoing land use changes and development. Importantly, red alder is also considered a bioindicator for ozone pollution and, like all plants, harbors a diverse endophyte community that may interact with aerial pollutants. In this study, we surveyed foliar fungal endophyte communities (microfungi) in red alder leaves from the metropolitan area of Portland, Oregon, USA, using culture-based techniques, and found that communities differed significantly by site. Our results suggest that the fungal endophyte community composition in red alder leaves may be influenced in part by local air pollution sources, likely in conjunction with other site characteristics. As urban areas expand, more studies should focus on how the urban environment affects plant–microbe community ecology and endophyte–host interactions, as well as on the long-term consequences for other ecosystem processes such as leaf litter decomposition.

Deficiencies of nutrients, especially phosphorus (P), and growing-season moisture may limit the growth and survival of nitrogen-fixing red alder (Alnus rubra Bong.), a commercially – important species in coastal forests of the Pacific Northwest and British Columbia. Future growing seasons are projected to be warmer and drier, potentially reducing P availability and plantation survival and growth on sites currently considered acceptable for alder management. We examined the effects of P additions on growth, survival, and nutrition in a young alder plantation on a moderately fertile and periodically droughty site on eastern Vancouver Island. Alder was planted in fall 1999 (C99 cohort) and fill-planted in spring 2001 (C01 cohort). We applied triple superphosphate (0-45-0) beginning in spring 2001 and repeatedly to year 6. Initial rates were 0, 15, or 30 g P tree−1 for the P0, P1, and P2 regimes respectively, and cumulative rates over 6 years were 0, 41, and 288 g P tree−1. Moisture availability was inferred from monthly precipitation data and periodic measurements of soil moisture content. Growth and mortality were measured yearly through year 13 and foliar elemental concentrations, yearly through year 6. Through 13 years, P-fertilized trees were taller (+1.6 m, +16%) than unfertilized trees, had greater diameter at breast height (dbh) (+1.5 cm; +14%) and, through year 6, greater foliar P concentrations. Treatment effects on growth were significant in year one and maintained thereafter but did not differ between the P1 and P2 regimes. Effects of P additions on current annual basal area increment (CAIba) increased with early growing-season precipitation. Total mortality was ca. 30%, two-fold greater in the fill-planted (C01) than in the C99 cohort, and greatest in dry summers (2003, 2004), but did not vary with P treatment. Effects of P additions on height and dbh were less for crop (largest-diameter 400 trees ha−1) than for non-crop trees. Through 13 years, P additions did not affect the diameter-density curve. Effects of P additions on growth were less through 3 years but greater through 8–10 years than in plantations on sites classified as moister and more fertile. P additions early in plantation development can increase the productivity of red alder in periodically dry sites, but the increases are greater during moister growing seasons.

Urban streams are impacted by invasion of exotic riparian plants and the accumulation of plastic trash, which alter in-stream litter subsidies, and cause changes that cascade up the aquatic food web. The impacts of these factors on urban streams is poorly understood. We compared decay rates and invertebrate colonizers of 5 litter pack types in 4 urban streams in Victoria, British Columbia, Canada: Native Red alder (Alnus rubra) and Sitka willow (Salix sitchensis), invasive English ivy (Hedera sp.), Himalayan blackberry (Rubus armeniacus) and plastic trash (i.e. Styrofoam (polystyrene (PS)), plastic bag (high-density polyethylene (HDPE)), and Mylar (polyethylene terephthalate (PET). We tested 4 hypotheses: 1) exotic ivy and blackberry leaves would decay more slowly than native leaves; 2) exotic ivy and blackberry leaves would attract fewer and less diverse stream invertebrates than native leaves; 3) plastic trash would decay more slowly than leaves; and, 4) plastic trash would attract fewer and less diverse stream invertebrates than leaves. We found no difference between the leaf litter decay rates, however plastic trash decayed more slowly than leaves. Trash decay rates were faster than reported in marine environments, suggesting that plastic trash removal should be a management priority. Stream invertebrates colonized all pack types equally. We observed significant differences in litter decay rates and invertebrate assemblage alpha and Shannon–Wiener diversities across the 4 streams - likely related to differences in stream-specific environmental attributes including flashiness, stream discharge, and biological decay. We conclude that site-specific decay forces supersede litter quality in Pacific Coast urban streams.

The invasion of the Pacific Northwest, USA by northern barred owls (Strix varia) is a conservation conundrum because it contributes to lingering declines in populations of northern spotted owls (Strix occidentalis caurina). We evaluated factors influencing nocturnal (i.e., foraging) habitat selection by northern barred owls using a repeated-studies design and information-theoretic methods across 3 Pacific Northwest study areas, each containing a broad range of forest and environmental conditions. We constructed discrete-choice resource selection functions (RSF) based upon telemetry points linked to forest inventory plots and map-based physical environmental metrics within home ranges of radio-tagged barred owls at Chehalis, Washington (n = 16), Springfield, Oregon (n = 22), and Arcata, California (n = 15). A general RSF based upon pooling data across study areas suggested barred owls selectively hunted for prey in lower-slope positions on southerly aspects often near streams at low elevations, and often within red alder (Alnus rubra) dominated stands or in moderately dense patches of medium- and large-diameter coniferous trees close to patches containing nests. The relative probability of use decreased with increasing densities of small-diameter trees, suggesting barred owls avoided clearcuts and young plantations. These general patterns were modified by study-area variation in tree species composition and density. Study-area-specific factors that were associated positively with barred owl habitat selection included increasing basal area of western redcedar (Thuja plicata) and red alder at Chehalis and increasing densities of western redcedar and basal area of bigleaf maple (Acer macrophyllum) and western hemlock (Tsuga heterophylla) at Springfield. At Arcata, high densities of Douglas-fir (Pseudotsuga menziesii) trees and increasing basal area of tanoak (Notholithocarpus densiflorus) were negatively associated with barred owl habitat selection. Seasonal patterns of habitat selection did not differ dramatically although model coefficients suggested selection for specific tree species was weaker in the non-breeding season and barred owls did not seek topographic situations that provided thermo-regulatory benefits. The information may help inform conservation strategies for reducing competition between barred owls and northern spotted owls or perhaps in predicting colonization of new areas by barred owls. © 2017 The Wildlife Society.

Regional and global studies of ectomycorrhizal fungal (EMF) communities associated with Alnus have made progress in determining the key factors that influence EMF diversity and biogeography. Smaller scale studies provide a valuable complement by relating EMF to environmental gradients and describing how community composition is influenced by competition and niche partitioning. We examined controls on EMF community composition in an 80 y-old Alnus rubra stand. EMF species were identified using root tip morphology and DNA sequencing, and related to soil variables (soil moisture, total C, total N, C:N ratio, PO4-P, soil pH) and to Frankia nodulation using Threshold Indicator Taxa ANalysis (TITAN) and multivariate techniques. Twenty-two EMF species were identified, including 14 that are new associates of A. rubra. EMF community composition varied temporally and was influenced by Frankia nodulation and soil chemistry. Species co-occurrence patterns suggest niche partitioning and competitive exclusion interact with subtle differences in the soil environment to influence EMF community composition.

In PNAS we report that the symbiotic nitrogen-fixing tree red alder ( Alnus rubra ) accesses more rock-derived nutrients than nonfixing trees (1). We further hypothesize that excess nitrogen fixation by alder, leading to high rates of nitrification and nitric acid generation in soil, could accelerate mineral weathering (1, 2). Krishna et al. (3) seek to provide an alternative explanation for accelerated mineral weathering by the closely related tree Himalayan alder ( Alnus nepalensis ). They invoke cluster roots and rhizosphere exudates as causes of enhanced mineral weathering (3) but provide no supporting evidence, which we have previously addressed (2). The remainder of their comment addresses age-related changes in nitrogen … [↵][1]1To whom correspondence may be addressed. Email: sperakis{at}usgs.gov. [1]: #xref-corresp-1-1

Timber durability is often assessed using small wood stakes exposed in direct soil contact, and the assessment generally emphasizes effects on wood rather than organisms involved. Understanding fungal colonization patterns can help identify key decay agents under varying conditions and use these patterns to improve wood protection strategies. Fungal colonization of red alder (Alnus rubra), Douglas-fir (Pseudotsuga menziesii) heartwood/sapwood, and western redcedar (Thuja plicata) field stakes was assessed over 2 y in western Oregon. Spatiotemporal fungal community variations were identified via culturing and DNA sequencing, where 814 isolates were identified from 84 stakes. Forty-six ascomycete genera were identified, with Phialophora, Trichoderma, and Epicoccum species occurring most frequently. Twenty-three basidiomycete genera were identified, with Trametes and Phanerochaete being the most common. Douglas-fir and western redcedar stakes contained the highest and lowest diversity levels, respectively, reflecting natural durability differences of these species. Fungal species abundance was higher below ground than in the above ground and groundline zones, likely reflecting more stable moisture regimes, proximity to soil-based fungi, and potential nutrient migration into wood beneath the soil surface. Ascomycetes were proportionally more abundant early in the exposure period, but basidiomycetes were also observed early in the process, and there appeared to be no consistent colonization pattern.

Symbiotic nitrogen (N)-fixing trees supply significant N inputs to forest ecosystems, leading to increased soil fertility, forest growth, and carbon storage. Rapid growth and stoichiometric constraints of N fixers also create high demands for rock-derived nutrients such as phosphorus (P), while excess fixed N can generate acidity and accelerate leaching of rock-derived nutrients such as calcium (Ca). This ability of N-fixing trees to accelerate cycles of Ca, P, and other rock-derived nutrients has fostered speculation of a direct link between N fixation and mineral weathering in terrestrial ecosystems. However, field evidence that N-fixing trees have enhanced access to rock-derived nutrients is lacking. Here we use strontium (Sr) isotopes as a tracer of nutrient sources in a mixed-species temperate rainforest to show that N-fixing trees access more rock-derived nutrients than nonfixing trees. The N-fixing tree red alder (Alnus rubra), on average, took up 8 to 18% more rock-derived Sr than five co-occurring nonfixing tree species, including two with high requirements for rock-derived nutrients. The increased access to rock-derived nutrients occurred despite spatial variation in community-wide Sr sources across the forest, and only N fixers had foliar Sr isotopes that differed significantly from soil exchangeable pools. We calculate that increased uptake of rock-derived nutrients by N-fixing alder requires a 64% increase in weathering supply of nutrients over nonfixing trees. These findings provide direct evidence that an N-fixing tree species can also accelerate nutrient inputs from rock weathering, thus increasing supplies of multiple nutrients that limit carbon uptake and storage in forest ecosystems.

Phytophthora alni, a soil- and waterborne pathogen, causes aggressive root and collar rot on riparian alder populations (1,2,4). The disease has been described from several European countries with a destructive impact in Great Britain (1,2). All European alder species and the red alder (Alnus rubra) are highly susceptible. P. alni has multiple variants that have been placed in three subspecies: P. alni subsp. alni, P. alni subsp. uniformis, and P. alni subsp. multiformis (1). In July 2009, a survey of symptoms of Phytophthora rot from A. glutinosa at 20 riparian stands along the Avia River in Galicia (northwest Spain) was conducted. Affected trees showed symptoms of Phytophthora rot including abnormally small, sparse, and yellowish foliage, dieback in the canopy, necroses of the inner bark and cambium, and bleeding cankers on the trunks (2,4). Phytophthora spp. were baited from saturated rhizosphere soil and watercourses using oak leaflets (4). Roots and tissue from fresh active inner bark lesions were transferred to selective medium V8-PARPH agar (4) and incubated for 7 days at 22°C in the dark. A Phytophthora sp. was isolated, transferred to carrot agar (CA), and incubated in the dark. Colonies were appressed, often irregular in outline, and with limited aerial mycelium (1). Growth on CA occurred from 4 to 31°C with optimum growth at 23 to 25°C. Chlamydospores were not observed. Ellipsoid, nonpapillate, noncaducous sporangia had a length/breadth average ratio of 1.4. Nesting and extended internal proliferation occurred. Oogonia, antheridia, and oospores were abundantly produced in a single culture. Oogonia with tapered stalks were spherical (mature oogonia 38 to 50 μm in diameter) and some had ornamented walls or bullate protuberances (1,2). Antheridia were large, amphigynous, and predominantly two-celled (23 to 37 × 16 to 23 μm). Oospores were plerotic. Distorted comma-shaped or smaller oogonia and aborted oospores were observed (1). Amplification of DNA was accomplished by using sequence-characterized amplification region-PCR primers (3). The amplicon sizes obtained were identical to P. alni subsp. alni (3). Internal transcribed spacer (ITS)-DNA and nadh1 mitochondrial gene were also amplified. DNA sequences of ITS and mt-DNA regions were deposited in GenBank (Nos. GU108602 and GU108603). Comparison of the sequences showed 100% homology with P. alni subsp. alni (GenBank Nos. FJ746679 and DQ202490). P. alni subsp. alni was recovered from trees at 3 of 20 riparian alder stands with symptoms. Pathogenicity of one representative isolate was confirmed by inoculating 10 3-year-old A. glutinosa seedlings grown in pots. One shallow cut was made into the bark at the collar level. A colonized agar plug, from the margin of an actively growing colony of P. alni subsp. alni, was inserted beneath the flap that was sealed with Parafilm. Five controls seedlings received only sterile CA agar plugs. Plants were incubated at 24°C and 95% humidity for 30 days. On inoculated plants, necroses progressed bidirectionally from the wound, and dead leaves and wilting of shoots were observed. P. alni subsp. alni was recovered from inoculated seedlings, but not from controls. To our knowledge, this is the first report of Phytophthora rot on alder caused by P. alni subsp. alni in Spain. References: (1) C. M. Brasier et al. Mycol. Res. 108:1172, 2004. (2) J. Gibbs et al. For. Comm. Bull. 126, 2003 (3) R. Ioos et al. Eur. J. Plant Pathol. 112:323, 2005. (4) T. Jung et al. Plant Pathol. 53:197, 2004.

The potential for limiting fungal attack on red alder (Alnus rubra Bong.) pallet stock was evaluated in a small-scale field test. Alder is extremely susceptible to biological attack, as evidenced by the nearly complete colonization of nontreated materials within 18 days after cutting. Pallet stock was either dipped or sprayed with 11 different candidate fungicides. Dipping tended to produce better protection than spraying, reflecting the opportunity for greater uptake during the dipping process. Most treatments provided protection for 11 days when applied by dipping, while only four treatments were able to provide protection for the full 30-day test period. The results suggest that chemical protection of alder is possible, but the protective period is much shorter than that found with other wood species.

Background Experiencing an adequate amount of cold temperatures over winter is necessary for many temperate tree species to break dormancy and flower in spring. Thus, changes in winter and spring temperatures associated with climate change may influence when trees break dormancy and flower in the future. There have been several experimental studies that have quantified the effectiveness of cold temperatures for chilling requirements for vegetative budburst of temperate trees; however, there are few experimental studies addressing the chilling requirements for reproductive budburst of trees, as it is difficult to place reproductively mature trees in temperature-controlled environments. Methods To identify how changing temperatures associated with climate change may impact reproductive phenology, we completed a temperature-controlled growth chamber experiment using cuttings of reproductive branches of red alder (Alnus rubra), one of the most widespread hardwood tree species of the Pacific Northwest, USA. The purpose of this study was to examine how colder (4 °C) and warmer (9 °C) winter temperature regimes influenced the timing of reproductive budburst of red alder cuttings in spring. We also compared the date of budburst of cuttings to that of branches from intact trees. Results We found that cuttings flowered earlier after pretreatment with a 4 °C winter temperature regime than after a 9 °C winter temperature regime. We found no significant differences between the timing of male budburst of cuttings exposed to ambient conditions compared to male budburst of branches from intact trees. We used our experimental data to estimate a “possibility-line” that shows the accumulated chilling and forcing temperatures necessary prior to reproductive budburst of red alder. Discussion This study provides a preliminary indication that warmer winters with climate change may not be as effective as colder winters for satisfying chilling temperature requirements of a Northwest hardwood tree species.

Perakis and Pett-Ridge (1) recently reported in PNAS that N2-fixing red alder ( Alnus rubra ) trees obtain significantly more rock-derived strontium than 5 codominant nonfixing trees in a mixed temperate rainforest in the Oregon Coast Range. The authors ascribe this to the fact that excess fixed N generates acidity, accelerating leaching of rock-derived nutrients. However, what the authors do not point out is that Alnus spp. produce cluster roots (2). In fact, actinorhizal species in different families produce cluster roots, and we are unaware of any actinorhizal species that does not produce cluster roots (3). Cluster roots mobilize poorly available phosphorus and other elements because they release large amounts of carboxylates (3). Functionally equivalent but morphologically very different specialized roots occur in, for example, Cactaceae (4), Cyperaceae (5), and Velloziaceae (6). Carboxylate-releasing cluster roots in … [↵][1]1To whom correspondence may be addressed. Email: hans.lambers{at}uwa.edu.au. [1]: #xref-corresp-1-1

The symbiotic nitrogen (N)-fixing tree red alder ( Alnus rubra ) is an ecologically and economically important species that has been studied for over 50 y (1, 2). Our recent finding that red alder can preferentially access rock-derived nutrients provides additional insight into the ecology of this N-fixing tree. Our manuscript evaluates several explanations for how red alder eats rocks and suggests that the main driver is nitric acid (3). Lambers et al. (4) agree that red alder preferentially accesses rock-derived nutrients, but instead hypothesize that red alder produces cluster roots, which mobilize rock-derived nutrients by releasing carboxylates. Our analysis builds on widespread observations that excess fixed N … [↵][1]1To whom correspondence should be addressed. Email: sperakis{at}usgs.gov. [1]: #xref-corresp-1-1

The White-footed Vole (Arborimus albipes) is one of the least-studied small mammals in North America. Most reported occurrence data come from incidental captures from larger trapping efforts and have demonstrated a strong affinity for Red Alder (Alnus rubra) trees. Recent research represented a range expansion in western Oregon, but little is known about the species at the southern end of its range in California. We developed a distribution model for the species to identify areas best targeted for additional White-footed Vole research. We used this model to survey by trapping in Humboldt and Del Norte Counties, California, and documented 3 new occurrences for the species. These 3 occurrences were incorporated into a final distribution model. This model suggests the possibility of White-footed Vole occurrence in southern Humboldt and northern Mendocino Counties, which would represent a southern expansion of their range. White-footed Voles were captured in areas with smaller alders than non-capture...

Organic fillers derived from biomass waste streams are commonly formulated with phenol-formaldehyde resoles in the production of veneer-based wood composites. The surface characteristics of three fillers, flours of walnut shell (Juglans regia), red alder (Alnus rubra) bark, and corn cob (Zea mays) furfural production residue, were investigated with the column wicking method using the Washburn equation. A series of n-alkanes (spreading liquids) provided reliable estimates of interstitial pore radii, which then enabled contact angle estimates for a variety of non-spreading liquids. Among the three fillers, only alder bark data were complicated by swelling effects, which were eliminated with strict data quality criteria. Corn cob residue exhibited the highest total surface free energy, but all fillers had low energies that were dominated by dispersive effects. The minor polar contributions were basic. Specific surface areas determined from wicking substantially disagreed with Brunauer–Emmett–Teller m...

Ammonia oxidation, the first step of nitrification, is mediated by both ammonia-oxidizing archaea (AOA) and bacteria (AOB); however, the relative contributions of AOA and AOB to soil nitrification are not well understood. In this study we used 1-octyne to discriminate between AOA- and AOB-supported nitrification determined both in soil-water slurries and in unsaturated whole soil at field moisture. Soils were collected from stands of red alder (Alnus rubra Bong.) and Douglas-fir (Pseudotsuga menziesii Mirb. Franco) at three sites (Cascade Head, the H.J. Andrews, and McDonald Forest) on acidic soils (pH 3.9–5.7) in Oregon, USA. The abundances of AOA and AOB were measured using quantitative PCR by targeting the amoA gene, which encodes subunit A of ammonia monooxygenase. Total and AOA-specific (octyne-resistant) nitrification activities in soil slurries were significantly higher at Cascade Head (the most acidic soils, pH

Studies were carried out to evaluate the suitability of Canadian prairie-grown, short-rotation hybrid poplar for the manufacture of high value solid wood products, including appearance grade lumber, plywood, laminated veneer lumber (LVL) and oriented strandboard (OSB). For appearance grade lumber applications, hybrid poplar was found to be an attractive wood as it machined and finished well. This wood can be finished to take on a number of different looks or to resemble other species. Hybrid poplar would appear suitable in applications such as millwork, mouldings, turnings and bedroom furniture where other softer hardwood species such as red alder (Alnus rubra Bong.) and yellow poplar (Liriodendron tulipifera L.) are currently being used. Hybrid poplar veneer was suitable for manufacturing high quality plywood, either used by itself, or mixed with other species such as aspen (Populus tremuloides Mich.) or spruce (Picea spp.). If the hybrid poplar veneer is stress graded, a portion of the veneer would be suitable for manufacturing structural grade LVL. Hybrid poplar can be processed commercially using procedures similar to those used for manufacturing aspen plywood and LVL. Properties of OSB manufactured from hybrid poplar were fully equivalent to those of OSB manufactured from aspen. Hybrid poplar can be substituted for aspen in OSB at any level with no apparent effect on panel properties.

Summary Described as one of the most destructive pathogens of agricultural crops and forest trees, Phytophthora is a genus of microorganisms containing over 100 known species. Phytophthora alni has caused collar and root disease in alders throughout Europe, and a subspecies has recently been isolated in North America. Reports of canopy dieback in red alder, Alnus rubra, prompted a survey of their overall health and to determine whether P. alni was present in western Oregon riparian ecosystems. Over 1100 Phytophthora isolates were recovered, representing 20 species and 2 taxa. Phytophthora-type cankers were observed in many trees, and their incidence was positively correlated with canopy dieback. High levels of mortality for red alder were not observed, which suggests these Phytophthora species are not aggressive pathogens. To test this hypothesis, three stem wound inoculations and one root dip were conducted on red alder seedlings using 13 Phytophthora species recovered from the riparian survey. Ten of the 13 Phytophthora species produced significant lesions in at least one pathogenicity test. Phytophthora siskiyouensis produced the largest lesions on red alder from the two stem wound inoculation tests conducted under summer conditions, while P. taxon Pgchlamydo caused the largest lesions during the winter stem wound inoculation test. Phytophthora gonapodyides, P. taxon Pgchlamydo and P. siskiyouensis have previously been found associated with necrotic alder roots and bole cankers in the field, and with the pathogenicity results reported here, we have established these species as causes of Phytophthora root disease and Phytophthora bole canker of alder in Oregon. While none of the Phytophthora species were especially aggressive towards red alder in the pathogenicity tests, they did cause localized disease symptoms. By weakening the root systems or boles of alders, the Phytophthoras could be leaving alders more susceptible to other insects and pathogens.

Summary 1. The phenology of many species is shifting in response to climatic changes, and these shifts are occurring at varying rates across species. This can potentially affect species’ interactions and individual fitness. However, few studies have experimentally tested the influence of warming on the timing of species interactions. This is an important gap in the literature given the potential for different direct and indirect effects of temperature via phenological change. 2. Our aim was to test the effects of warming on the western tent caterpillar (Malacosoma californicum pluviale). In addition to the direct effects of warming, we considered the two primary indirect effects mediated by warming-driven changes in its host plant, red alder (Alnus rubra): changes in resource availability due to phenological mismatch (i.e. changes in the relative timing of the interaction), and changes in resource quality associated with leaf maturation. 3. We experimentally warmed egg masses and larvae of the western tent caterpillar placed on branches of red alder in the field. 4. Warming advanced the timing of larval but not leaf emergence. This led to varying degrees of phenological mismatch, with larvae emerging as much as 25 days before to 10 days after the emergence of leaves. Even the earliest-emerging larvae, however, had high survival in the absence of leaves for up to 3 weeks, and they were surprisingly resistant to starvation. In addition, although warming created phenological mismatch that initially slowed the development of larvae that emerged before leaf emergence, it accelerated larval development once leaves were available. Therefore, warming had no net effect on our measures of insect performance. 5. Our results demonstrate that the indirect effects of warming, in creating phenological mismatch, are as important to consider as the direct effects on insect performance. Although future climatic warming might influence plants and insects in different ways, some insects may be well adapted to variation in the timing of their interactions.

We gathered baseline data to assess alder tree damage in western Oregon riparian ecosystems. We sought to determine if Phytophthora-type cankers found in Europe or the pathogen Phytophthora alni subsp. alni, which represent a major threat to alder forests in the Pacific Northwest, were present in the study area. Damage was evaluated in 88 transects; information was recorded on damage type (pathogen, insect or wound) and damage location. We evaluated 1445 red alder (Alnus rubra), 682 white alder (Alnus rhombifolia) and 181 thinleaf alder (Alnus incana spp. tenuifolia) trees. We tested the correlation between canopy dieback and canker symptoms because canopy dieback is an important symptom of Phytophthora disease of alder in Europe. We calculated the odds that alder canopy dieback was associated with Phytophthora-type cankers or other biotic cankers. P. alni subsp. alni (the causal agent of alder disease in Europe) was not identified in western Oregon; however, Phytophthora siskiyouensis was isolated from Phytophthora-type cankers which were present on 2% of red alder trees and 3% of white alder trees. The odds of canopy dieback were 5.4 and 4.8 times greater for red and white alder (respectively) with Phytophthora-type canker symptoms than in trees without such cankers. The percentage of trees with canopy dieback was 51%, 32%, and 10% for red, white, and thinleaf alder respectively. Other common damage included wounding, foliar pathogens and insects on red alder. This is the first report of Phytophthora canker of alder in United States forests and first report of P. siskiyouensis isolation from alder in forests anywhere.