David C. Shaw

Disturbances and structural development of natural forest ecosystems with silvicultural implications, using Douglas-fir forests as an example

Forest managers need a comprehensive scientific understanding of natural stand development processes when designing silvicultural systems that integrate ecological and economic objectives, including a better appreciation of the nature of disturbance regimes and the biological legacies, such as live trees, snags, and logs, that they leave behind. Most conceptual forest development models do not incorporate current knowledge of the: (1) complexity of structures (including spatial patterns) and developmental processes; (2) duration of development in long-lived forests; (3) complex spatial patterns of stands that develop in later stages of seres; and particularly (4) the role of disturbances in creating structural legacies that become key elements of the post-disturbance stands. We elaborate on existing models for stand structural development using natural stand development of the Douglas-fir—western hemlock sere in the Pacific Northwest as our primary example; most of the principles are broadly applicable while some processes (e.g. role of epicormic branches) are related to specific species. We discuss the use of principles from disturbance ecology and natural stand development to create silvicultural approaches that are more aligned with natural processes. Such approaches provide for a greater abundance of standing dead and down wood and large old trees, perhaps reducing short-term commercial productivity but ultimately enhancing wildlife habitat, biodiversity, and ecosystem function, including soil protection and nutrient retention. # 2002 Elsevier Science B.V. All rights reserved.

Epiphyte Habitats in an Old Conifer Forest in Western Washington, U.S.A.

Abstract Old conifer forests in the Pacific Northwest have a wide range of microhabitats induced by canopy structure and substrate characteristics. We used the Wind River Canopy Crane to sample lichens and bryophytes throughout the spectrum of habitats available to epiphytes. Of the 111 species found in 72 sample units, 97 were lichens and 14 were bryophytes. Epiphyte communities showed marked variation with respect to height in the canopy, bark vs. wood, degree of sheltering, and stem diameter. Of these factors, height in the canopy was most strongly related to epiphyte communities. Furthermore, the top two meters of the tallest trees hosted a diverse assemblage of both rare species (Tholurna dissimilis) and weedy, nitrophilous species (Candelaria concolor, Hypogymnia tubulosa, Parmelia sulcata), presumably induced by birds delivering lichen propagules and nutrients. Ten species were more frequent on bare wood than bark, including Ophioparma rubricosa, Letharia vulpina, Placynthiella spp., Ptychographa xylographoides, Trapeliopsis flexuosa, and Xylographa parallela. Species richness was highly variable, even within habitats. The only factor found related to species richness was height in the canopy, the middle and upper layers each having about twice the species per sample unit as lower in the canopy.

Comparison of dwarf mistletoes (Arceuthobium spp., Viscaceae) in the western United States with mistletoes (Amyema spp., Loranthaceae) in Australia: ecological analogs and reciprocal models for ecosystem management

Whereas the biology, physiology and systematics of mistletoes have been explored in considerable detail, their ecology has received less attention and our understanding is highly fragmentary. A conspicuous exception is the dwarf mistletoes (Arceuthobium spp.)—a genus that exclusively parasitises coniferous trees, including many commercially valuable species in the forests of the western United States. Accordingly, these plants have been the subjects of intensive cross-disciplinary research for the past five decades, initially from a control and management perspective but extending into most aspects of their ecology and life history. This review summarises our understanding of dwarf mistletoes, focusing on recent developments in the areas of mistletoe–wildlife interactions, fire, ecosystem ecology and conservation biology. We also compare dwarf mistletoes with Australian mistletoes in the genus Amyema, a diverse suite of species found throughout the continent. Despite fundamental differences in their evolutionary origin and most aspects of their autecology and life history, the genera exhibit many similarities in terms of their ecological role in forests and woodlands, and their influence on stand- and forest-scale dynamics. In particular, both groups provide nesting resources for a range of birds and mammals, and nutritional resources for a diverse assemblage of species. Both also interact with fire, potentially leading to changes in successional dynamics at the stand scale. At an applied level, both groups are widely considered as pests but, as our understanding of these keystone species improves, they have the potential to serve as sensitive ecological indicators for their respective ecosystems. Key research priorities are identified for further research on both groups of mistletoes and more explicit comparative research, with Arceuthobium serving as a valuable template for future work on Amyema and Australian mistletoes in general.

Crown structure and the distribution of epiphyte functional group biomass in old-growth Pseudotsuga menziesii trees

Epiphyte functional groups (alectorioid lichens, cyanolichens, other lichens, and bryophytes) were sampled in nine old-growth, canopy-emergent, Pseudotsuga menziesii trees along a riparian corridor...

A review of logistic regression models used to predict post-fire tree mortality of western North American conifers

Logistic regression models used to predict tree mortality are critical to post-fire management, planning prescribed burns and understanding disturbance ecology. We review literature concerning post-fire mortality prediction using logistic regression models for coniferous tree species in the western USA. We include synthesis and review of: methods to develop, evaluate and interpret logistic regression models; explanatory variables in logistic regression models; factors influencing scope of inference and model limitations; model validation; and management applications. Logistic regression is currently the most widely used and available technique for predicting post-fire tree mortality. Over 100 logistic regression models have been developed to predict post-fire tree mortality for 19 coniferous species following wild and prescribed fires. The most widely used explanatory variables in post-fire tree mortality logistic regression models have been measurements of crown (e.g. crown scorch) and stem (e.g. bole char) injury. Prediction of post-fire tree mortality improves when crown and stem variables are used collectively. Logistic regression models that predict post-fire tree mortality are the basis of simple field tools and contribute to larger fire-effects models. Future post-fire tree mortality prediction models should include consistent definition of model variables, model validation and direct incorporation of physiological responses that link to process modelling efforts.

Does wildfire likelihood increase following insect outbreaks in conifer forests

Although there is acute concern that insect-caused tree mortality increases the likelihood or severity of subsequent wildfire, previous studies have been mixed, with findings typically based on stand-scale simulations or individual events. This study investigates landscape- and regional-scale wildfire likelihood following outbreaks of the two most prevalent native insect pests in the US Pacific Northwest (PNW): mountain pine beetle (MPB; Dendroctonus ponderosae) and western spruce budworm (WSB; Choristoneura freemani). We leverage seamless census data across numerous insect and fire events to (1) summarize the interannual dynamics of insects (1970–2012) and wildfires (1984–2012) across forested ecoregions of the PNW; (2) identify potential linked disturbance interactions with an empirical wildfire likelihood index; (3) quantify this insect-fire likelihood across different insect agents, time lags, ecoregions, and fire sizes. All three disturbance agents have occurred primarily in the drier, interior conif...

Seasonal carbohydrate dynamics and growth in Douglas-fir trees experiencing chronic, fungal-mediated reduction in functional leaf area

Stored non-structural carbohydrates (NSCs) could play an important role in tree survival in the face of a changing climate and associated stress-related mortality. We explored the effects of the stomata-blocking and defoliating fungal disease called Swiss needle cast on Douglas-fir carbohydrate reserves and growth to evaluate the extent to which NSCs can be mobilized under natural conditions of low water stress and restricted carbon supply in relation to potential demands for growth. We analyzed the concentrations of starch, sucrose, glucose and fructose in foliage, twig wood and trunk sapwood of 15 co-occurring Douglas-fir trees expressing a gradient of Swiss needle cast symptom severity quantified as previous-year functional foliage mass. Growth (mean basal area increment, BAI) decreased by ∼80% and trunk NSC concentration decreased by 60% with decreasing functional foliage mass. The ratio of relative changes in NSC concentration and BAI, an index of the relative priority of storage versus growth, more than doubled with increasing disease severity. In contrast, twig and foliage NSC concentrations remained nearly constant with decreasing functional foliage mass. These results suggest that under disease-induced reductions in carbon supply, Douglas-fir trees retain NSCs (either actively or due to sequestration) at the expense of trunk radial growth. The crown retains the highest concentrations of NSC, presumably to maintain foliage growth and shoot extension in the spring, partially compensating for rapid foliage loss in the summer and fall.

STAND-LEVEL HERBIVORY IN AN OLD-GROWTH CONIFER FOREST CANOPY

Abstract Herbivory is an important ecological process in forest canopies but is difficult to measure, especially for whole stands. We used the Wind River Canopy Crane in Washington State to access 101 randomly-located sample points throughout the forest canopy. This provided a relatively quick and convenient way to estimate herbivory for a whole stand. The overall level of herbivory was estimated at 1.6% of leaf area. The distribution was strongly skewed to the lower canopy where broad-leafed species experienced higher levels of herbivory. Herbivory averaged 0.3% in conifers and 13.5% in broad-leafed species. Fully half of the sample points had no detectable herbivory. Herbivory in this old-growth conifer forest is among the lowest levels published for forests around the globe and may reflect the general levels of herbivory in temperate coniferous forests during nonoutbreak conditions. Our whole-stand estimate is the first attempt at measuring herbivory for an entire forest stand in the Pacific Northwest.

Tree-ring stable isotopes record the impact of a foliar fungal pathogen on CO(2) assimilation and growth in Douglas-fir.

Swiss needle cast (SNC) is a fungal disease of Douglas-fir (Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree-rings of Douglas-fir and a non-susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ(13)C) of treated Douglas-fir tree-rings was greater than that of untreated Douglas-fir tree-rings during the fungicide treatment period. Both annual growth and tree-ring Δ(13)C increased with treatment such that treated Douglas-fir had values similar to co-occurring western hemlock during the treatment period. There was no difference in the tree-ring oxygen stable isotope ratio between treated and untreated Douglas-fir. Tree-ring Δ(13)C of diseased Douglas-fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years.

Impacts of dwarf mistletoe on the physiology of host Tsuga heterophylla trees as recorded in tree-ring C and O stable isotopes

Dwarf mistletoes, obligate, parasitic plants with diminutive aerial shoots, have long-term effects on host tree water relations, hydraulic architecture and photosynthetic gas exchange and can eventually induce tree death. To investigate the long-term (1886-2010) impacts of dwarf mistletoe on the growth and gas exchange characteristics of host western hemlock, we compared the diameter growth and tree-ring cellulose stable carbon (C) and oxygen (O) isotope ratios (δ(13)Ccell, δ(18)Ocell) of heavily infected and uninfected trees. The relative basal area growth of infected trees was significantly greater than that of uninfected trees in 1886-90, but declined more rapidly in infected than uninfected trees through time and became significantly lower in infected than uninfected trees in 2006-10. Infected trees had significantly lower δ(13)Ccell and δ(18)Ocell than uninfected trees. Differences in δ(18)Ocell between infected and uninfected trees were unexpected given that stomatal conductance and environmental variables that were expected to influence the δ(18)O values of leaf water were similar for both groups. However, estimates of mesophyll conductance (gm) were significantly lower and estimates of effective path length for water movement (L) were significantly higher in leaves of infected trees, consistent with their lower values of δ(18)Ocell. This study reconstructs the long-term physiological responses of western hemlock to dwarf mistletoe infection. The long-term diameter growth and δ(13)Ccell trajectories suggested that infected trees were growing faster than uninfected trees prior to becoming infected and subsequently declined in growth and leaf-level photosynthetic capacity compared with uninfected trees as the dwarf mistletoe infection became severe. This study further points to limitations of the dual-isotope approach for identifying sources of variation in δ(13)Ccell and indicates that changes in leaf internal properties such as gm and L that affect δ(18)Ocell must be considered.