Mark E. Swanson

The forgotten stage of forest succession: early‐successional ecosystems on forest sites

Early-successional forest ecosystems that develop after stand-replacing or partial disturbances are diverse in species, processes, and structure. Post-disturbance ecosystems are also often rich in biological legacies, including surviving organisms and organically derived structures, such as woody debris. These legacies and post-disturbance plant communities provide resources that attract and sustain high species diversity, including numerous early-successional obligates, such as certain woodpeckers and arthropods. Early succession is the only period when tree canopies do not dominate the forest site, and so this stage can be characterized by high productivity of plant species (including herbs and shrubs), complex food webs, large nutrient fluxes, and high structural and spatial complexity. Different disturbances contrast markedly in terms of biological legacies, and this will influence the resultant physical and biological conditions, thus affecting successional pathways. Management activities, such as post-disturbance logging and dense tree planting, can reduce the richness within and the duration of early-successional ecosystems. Where maintenance of biodiversity is an objective, the importance and value of these natural early-successional ecosystems are underappreciated.

Ecological importance of large-diameter trees in a temperate mixed-conifer forest.

Large-diameter trees dominate the structure, dynamics and function of many temperate and tropical forests. Although both scaling theory and competition theory make predictions about the relative composition and spatial patterns of large-diameter trees compared to smaller diameter trees, these predictions are rarely tested. We established a 25.6 ha permanent plot within which we tagged and mapped all trees ≥1 cm dbh, all snags ≥10 cm dbh, and all shrub patches ≥2 m2. We sampled downed woody debris, litter, and duff with line intercept transects. Aboveground live biomass of the 23 woody species was 507.9 Mg/ha, of which 503.8 Mg/ha was trees (SD = 114.3 Mg/ha) and 4.1 Mg/ha was shrubs. Aboveground live and dead biomass was 652.0 Mg/ha. Large-diameter trees comprised 1.4% of individuals but 49.4% of biomass, with biomass dominated by Abies concolor and Pinus lambertiana (93.0% of tree biomass). The large-diameter component dominated the biomass of snags (59.5%) and contributed significantly to that of woody debris (36.6%). Traditional scaling theory was not a good model for either the relationship between tree radii and tree abundance or tree biomass. Spatial patterning of large-diameter trees of the three most abundant species differed from that of small-diameter conspecifics. For A. concolor and P. lambertiana, as well as all trees pooled, large-diameter and small-diameter trees were spatially segregated through inter-tree distances <10 m. Competition alone was insufficient to explain the spatial patterns of large-diameter trees and spatial relationships between large-diameter and small-diameter trees. Long-term observations may reveal regulation of forest biomass and spatial structure by fire, wind, pathogens, and insects in Sierra Nevada mixed-conifer forests. Sustaining ecosystem functions such as carbon storage or provision of specialist species habitat will likely require different management strategies when the functions are performed primarily by a few large trees as opposed to many smaller trees.

Spatially nonrandom tree mortality and ingrowth maintain equilibrium pattern in an old‐growth Pseudotsuga–Tsuga forest

Mortality processes in old-growth forests are generally assumed to be driven by gap-scale disturbance, with only a limited role ascribed to density-dependent mortality, but these assumptions are rarely tested with data sets incorporating repeated measurements. Using a 12-ha spatially explicit plot censused 13 years apart in an approximately 500-year-old Pseudotsuga-Tsuga forest, we demonstrate significant density-dependent mortality and spatially aggregated tree recruitment. However, the combined effect of these strongly nonrandom demographic processes was to maintain tree patterns in a state of dynamic equilibrium. Density-dependent mortality was most pronounced for the dominant late-successional species, Tsuga heterophylla. The long-lived, early-seral Pseudotsuga menziesii experienced an annual stem mortality rate of 0.84% and no new recruitment. Late-seral species Tsuga and Abies amabilis had nearly balanced demographic rates of ingrowth and mortality. The 2.34% mortality rate for Taxus brevifolia was higher than expected, notably less than ingrowth, and strongly affected by proximity to Tsuga. Large-diameter Tsuga structured both the regenerating conspecific and heterospecific cohorts with recruitment of Tsuga and Abies unlikely in neighborhoods crowded with large-diameter competitors (P < 0.001). Density-dependent competitive interactions strongly shape forest communities even five centuries after stand initiation, underscoring the dynamic nature of even equilibrial old-growth forests.

The Importance of Large-Diameter Trees to Forest Structural Heterogeneity

Large-diameter trees dominate the structure, dynamics and function of many temperate and tropical forests. However, their attendant contributions to forest heterogeneity are rarely addressed. We established the Wind River Forest Dynamics Plot, a 25.6 ha permanent plot within which we tagged and mapped all 30,973 woody stems ≥1 cm dbh, all 1,966 snags ≥10 cm dbh, and all shrub patches ≥2 m2. Basal area of the 26 woody species was 62.18 m2/ha, of which 61.60 m2/ha was trees and 0.58 m2/ha was tall shrubs. Large-diameter trees (≥100 cm dbh) comprised 1.5% of stems, 31.8% of basal area, and 17.6% of the heterogeneity of basal area, with basal area dominated by Tsuga heterophylla and Pseudotsuga menziesii. Small-diameter subpopulations of Pseudotsuga menziesii, Tsuga heterophylla and Thuja plicata, as well as all tree species combined, exhibited significant aggregation relative to the null model of complete spatial randomness (CSR) up to 9 m (P≤0.001). Patterns of large-diameter trees were either not different from CSR (Tsuga heterophylla), or exhibited slight aggregation (Pseudotsuga menziesii and Thuja plicata). Significant spatial repulsion between large-diameter and small-diameter Tsuga heterophylla suggests that large-diameter Tsuga heterophylla function as organizers of tree demography over decadal timescales through competitive interactions. Comparison among two forest dynamics plots suggests that forest structural diversity responds to intermediate-scale environmental heterogeneity and disturbances, similar to hypotheses about patterns of species richness, and richness- ecosystem function. Large mapped plots with detailed within-plot environmental spatial covariates will be required to test these hypotheses.

Spatial aspects of tree mortality strongly differ between young and old‐growth forests

Rates and spatial patterns of tree mortality are predicted to change during forest structural development. In young forests, mortality should be primarily density dependent due to competition for light, leading to an increasingly spatially uniform pattern of surviving trees. In contrast, mortality in old-growth forests should be primarily caused by contagious and spatially autocorrelated agents (e.g., insects, wind), causing spatial aggregation of surviving trees to increase through time. We tested these predictions by contrasting a three-decade record of tree mortality from replicated mapped permanent plots located in young (< 60-year-old) and old-growth (> 300-year-old) Abies amabilis forests. Trees in young forests died at a rate of 4.42% per year, whereas trees in old-growth forests died at 0.60% per year. Tree mortality in young forests was significantly aggregated, strongly density dependent, and caused live tree patterns to become more uniform through time. Mortality in old-growth forests was spatially aggregated, but was density independent and did not change the spatial pattern of surviving trees. These results extend current theory by demonstrating that density-dependent competitive mortality leading to increasingly uniform tree spacing in young forests ultimately transitions late in succession to a more diverse tree mortality regime that maintains spatial heterogeneity through time.

Effects of remedial sport hunting on cougar complaints and livestock depredations.

Remedial sport hunting of predators is often used to reduce predator populations and associated complaints and livestock depredations. We assessed the effects of remedial sport hunting on reducing cougar complaints and livestock depredations in Washington from 2005 to 2010 (6 years). The number of complaints, livestock depredations, cougars harvested, estimated cougar populations, human population and livestock populations were calculated for all 39 counties and 136 GMUs (game management units) in Washington. The data was then analyzed using a negative binomial generalized linear model to test for the expected negative relationship between the number of complaints and depredations in the current year with the number of cougars harvested the previous year. As expected, we found that complaints and depredations were positively associated with human population, livestock population, and cougar population. However, contrary to expectations we found that complaints and depredations were most strongly associated with cougars harvested the previous year. The odds of increased complaints and livestock depredations increased dramatically (36 to 240%) with increased cougar harvest. We suggest that increased young male immigration, social disruption of cougar populations, and associated changes in space use by cougars - caused by increased hunting resulted in the increased complaints and livestock depredations. Widespread indiscriminate hunting does not appear to be an effective preventative and remedial method for reducing predator complaints and livestock depredations.

Identifying best methods for routine ELISA detection of microcystin in seafood

Ingestion of water contaminated with the cyanotoxin, microcystin (MC), can pose serious health risks to humans. MC is also known to accumulate in seafood; however, this exposure pathway is much less understood. A fundamental element of this uncertainty is related to analytical difficulties. Commercially available enzyme-linked immunosorbent assays (ELISAs) offer one of the best options for routine MC detection, but methods of detecting MC in tissue are far from standardized. We spiked freshwater finfish and marine mussel tissues with MC, then compared recovery rates using four different preparation protocols and two ELISA types (polyclonal anti-MC-ADDA/direct monoclonal (DM)). Preparation protocol, type of ELISA, and seafood tissue variety significantly affected MC detection. This is the first known study to use DM ELISA for tissue analyses, and our findings demonstrate that DM ELISA combined with a short solvent extraction results in fewer false positives than other commonly used methods. This method can be used for rapid and reliable MC detection in seafood.

Newman Lake restoration: A case study Part III. Hypolimnetic oxygenation

Abstract Since the mid-20th century, oxygen status at the sediment–water interface (SWI) has been implicated in regulating lake internal phosphorus loading. In deeper lakes, summer hypolimnetic oxygen depletion may thus play a critical role in phosphorus cycling and lake trophic status. Hypolimnetic aeration (HA) has been utilized for more than 4 decades to prevent development of anoxia, decrease internal phosphorus load, and enhance fisheries. Most recently, interest has shifted to hypolimnetic oxygenation (HO) for potential performance and economic advantages. In Newman Lake, hypolimnetic oxygenation was initiated in 1992 when the first lake application of downflow contact bubble oxygenation (Speece Cone technology) was installed. Oxygenation at Newman has reduced growing season Nürnberg Anoxic Factors (AF) from a range of about 30–60 d to <10 d. We propose that differences in predicted versus observed AF based on phosphorus may be utilized to assess lake restoration performance. Newman Lake has demonstrated the importance of operating the system at full capacity, as lower oxygen delivery rates do not produce proportional hypolimnetic oxygen concentrations, as well as other insights into HO system sizing and design.

Evaluating Integrated Watershed Management using multiple criteria analysis—a case study at Chittagong Hill Tracts in Bangladesh

Criteria and indicators assessment is one of the ways to evaluate management strategies for mountain watersheds. One framework for this, Integrated Watershed Management (IWM), was employed at Chittagong Hill Tracts region of Bangladesh using a multi-criteria analysis approach. The IWM framework, consisting of the design and application of principles, criteria, indicators, and verifiers (PCIV), facilitates active participation by diverse professionals, experts, and interest groups in watershed management, to explicitly address the demands and problems to measure the complexity of problems in a transparent and understandable way. Management alternatives are developed to fulfill every key component of IWM considering the developed PCIV set and current situation of the study area. Different management strategies, each focusing on a different approach (biodiversity conservation, flood control, soil and water quality conservation, indigenous knowledge conservation, income generation, watershed conservation, and landscape conservation) were assessed qualitatively on their potential to improve the current situation according to each verifier of the criteria and indicator set. Analytic Hierarchy Process (AHP), including sensitivity analysis, was employed to identify an appropriate management strategy according to overall priorities (i.e., different weights of each principle) of key informants. The AHP process indicated that a strategy focused on conservation of biodiversity provided the best option to address watershed-related challenges in the Chittagong Hill Tracts, Bangladesh.

Community composition and allometry of Leucothoe davisiae, Cornus sericea, and Chrysolepis sempervirens

Leucothoe davisiae Torr. ex A. Gray (Ericaceae), Sierra laurel, is an endemic shrub of the Sierra Nevada, Klamath Mountains, and Warner Mountains. We compared the woody plant community and allometry of Leucothoe with one widespread wetland shrub, red osier dogwood (Cornus sericea L. (Cornaceae)), and one widespread upland shrub, bush chinquapin (Chrysolepis sempervirens (Kellogg) Hjelmq. (Fagaceae)), in an Abies concolor – Pinus lambertiana forest. We examined 2282 mapped shrub patches and 34 392 trees in the Yosemite Forest Dynamics Plot (25.6 ha), Yosemite National Park, California, USA. We dissected 40–41 stems of each shrub species and determined foliage, bark, wood, and total biomass as a function of stem diameter. Community compositions determined from plants within a 5 m buffer of Leucothoe were different from those reported for Leucothoe communities in the Siskiyou National Forest and the Central Klamath. Leucothoe allometry based on basal diameter was more variable (total biomass R2 = 0.64; P < 0...