James A. Lutz

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 (SDu200a=u200a114.3 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.

Scale‐dependent relationships between tree species richness and ecosystem function in forests

1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity.

The Science of Firescapes: Achieving Fire-Resilient Communities

Abstract Wildland fire management has reached a crossroads. Current perspectives are not capable of answering interdisciplinary adaptation and mitigation challenges posed by increases in wildfire risk to human populations and the need to reintegrate fire as a vital landscape process. Fire science has been, and continues to be, performed in isolated “silos,” including institutions (e.g., agencies versus universities), organizational structures (e.g., federal agency mandates versus local and state procedures for responding to fire), and research foci (e.g., physical science, natural science, and social science). These silos tend to promote research, management, and policy that focus only on targeted aspects of the “wicked” wildfire problem. In this article, we provide guiding principles to bridge diverse fire science efforts to advance an integrated agenda of wildfire research that can help overcome disciplinary silos and provide insight on how to build fire-resilient communities.

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.

Plant diversity increases with the strength of negative density dependence at the global scale

Maintaining tree diversity Negative interaction among plant species is known as conspecific negative density dependence (CNDD). This ecological pattern is thought to maintain higher species diversity in the tropics. LaManna et al. tested this hypothesis by comparing how tree species diversity changes with the intensity of local biotic interactions in tropical and temperate latitudes (see the Perspective by Comita). Stronger local specialized biotic interactions seem to prevent erosion of biodiversity in tropical forests, not only by limiting populations of common species, but also by strongly stabilizing populations of rare species, which tend to show higher CNDD in the tropics. Science, this issue p. 1389; see also p. 1328 A global analysis of ~3000 species and ~2.4 million trees elucidates variations in tree species diversity between tropical and temperate latitudes. Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.

Comparative evolutionary diversity and phylogenetic structure across multiple forest dynamics plots: a mega-phylogeny approach

Forest dynamics plots, which now span longitudes, latitudes, and habitat types across the globe, offer unparalleled insights into the ecological and evolutionary processes that determine how species are assembled into communities. Understanding phylogenetic relationships among species in a community has become an important component of assessing assembly processes. However, the application of evolutionary information to questions in community ecology has been limited in large part by the lack of accurate estimates of phylogenetic relationships among individual species found within communities, and is particularly limiting in comparisons between communities. Therefore, streamlining and maximizing the information content of these community phylogenies is a priority. To test the viability and advantage of a multi-community phylogeny, we constructed a multi-plot mega-phylogeny of 1347 species of trees across 15 forest dynamics plots in the ForestGEO network using DNA barcode sequence data (rbcL, matK, and psbA-trnH) and compared community phylogenies for each individual plot with respect to support for topology and branch lengths, which affect evolutionary inference of community processes. The levels of taxonomic differentiation across the phylogeny were examined by quantifying the frequency of resolved nodes throughout. In addition, three phylogenetic distance (PD) metrics that are commonly used to infer assembly processes were estimated for each plot [PD, Mean Phylogenetic Distance (MPD), and Mean Nearest Taxon Distance (MNTD)]. Lastly, we examine the partitioning of phylogenetic diversity among community plots through quantification of inter-community MPD and MNTD. Overall, evolutionary relationships were highly resolved across the DNA barcode-based mega-phylogeny, and phylogenetic resolution for each community plot was improved when estimated within the context of the mega-phylogeny. Likewise, when compared with phylogenies for individual plots, estimates of phylogenetic diversity in the mega-phylogeny were more consistent, thereby removing a potential source of bias at the plot-level, and demonstrating the value of assessing phylogenetic relationships simultaneously within a mega-phylogeny. An unexpected result of the comparisons among plots based on the mega-phylogeny was that the communities in the ForestGEO plots in general appear to be assemblages of more closely related species than expected by chance, and that differentiation among communities is very low, suggesting deep floristic connections among communities and new avenues for future analyses in community ecology.

Towards a new paradigm in fire severity research using dose–response experiments

Most landscape-scale fire severity research relies on correlations between field measures of fire effects and relatively simple spectral reflectance indices that are not direct measures of heat output or changes in plant physiology. Although many authors have highlighted limitations of this approach and called for improved assessments of severity, others have suggested that the operational utility of such a simple approach makes it acceptable. An alternative pathway to evaluate fire severity that bridges fire combustion dynamics and ecophysiology via dose–response experiments is presented. We provide an illustrative example from a controlled nursery combustion laboratory experiment. In this example, severity is defined through changes in the ability of the plant to assimilate carbon at the leaf level. We also explore changes in the Differenced Normalised Differenced Vegetation Index (dNDVI) and the Differenced Normalised Burn Ratio (dNBR) as intermediate spectral indices. We demonstrate the potential of this methodology and propose dose–response metrics for quantifying severity in terms of carbon cycle processes.

The Evolution of Long-Term Data for Forestry: Large Temperate Research Plots in an Era of Global Change

Abstract When forest ecosystems develop over millennia, trees live five centuries, and mortality unfolds over decades, direct repeated observation (hereafter, longitudinal data) may be the only way to understand the fate of forests. Longitudinal data sets contribute greatly to our understanding, complementing experimental, modeling, and chronosequence approaches. Changing climate is changing forests, perhaps most rapidly through altered mortality regimes, and the elusive nature of integrated mechanistic understanding requires refinements and extensions to historically productive protocols. Changing climate reduces the inferential power of chronosequence techniques and changes model parameterization, and only some of the different factors contributing to tree mortality are expected to respond to climate variability and change. Because annual tree mortality rates are 5% to 1% for trees ≥ 1 cm dbh in young and old forests (respectively), detecting changes in mortality rates requires tracking thousands of trees, particularly to examine rare sub-populations of concern (e.g., large-diameter trees). And because mortality factors can be spatially aggregated and density-dependent, the causes and rates of tree mortality depend on the relationships between forest spatial structure and the direct and indirect effects of climate. Permanent plots with a combination of larger size, higher spatial precision, and greater sampling frequency will be required to further elucidate spatially explicit aspects of western forest demography. The combination of the longitudinal protocols developed by the Smithsonian Center for Tropical Forest Science, originally for studying tropical forest species diversity, and those developed by the US Geological Survey for annual tree mortality assessment together uniquely allow robust investigation of climate-mediated change in temperate forests.

Evaluating a new method for reconstructing forest conditions from General Land Office survey records

Historical forest conditions are often used to inform contemporary management goals because historical forests are considered to be resilient to ecological disturbances. The General Land Office (GLO) surveys of the late 19th and early 20th centuries provide regionally quasi-contiguous data sets of historical forests across much of the Western United States. Multiple methods exist for estimating tree density from point-based sampling such as the GLO surveys, including distance-based and area-based approaches. Area-based approaches have been applied in California mixed-conifer forests but their estimates have not been validated. To assess the accuracy and precision of plotless density estimators with potential for application to GLO data in this region, we imposed a GLO sampling scheme on six mapped forest stands of known densities (159-784 trees/ha) in the Sierra Nevada in California, USA, and Baja California Norte, Mexico. We compared three distance-based plotless density estimators (Cottam, Pollard, and Morisita) as well as two Voronoi area (VA) estimators, the Delincé and mean harmonic Voronoi density (MHVD), to the true densities. We simulated sampling schemes of increasing intensity to assess sampling error. The relative error (RE) of density estimates for the GLO sampling scheme ranged from 0.36 to 4.78. The least biased estimate of tree density in every stand was obtained with the Morisita estimator and the most biased was obtained with the MHVD estimator. The MHVD estimates of tree density were 1.2-3.8 times larger than the true densities and performed best in stands subject to fire exclusion for 100xa0yr. The Delincé approach obtained accurate estimates of density, implying that the Voronoi approach is theoretically sound but that its application in the MHVD was flawed. The misapplication was attributed to two causes: (1) the use of a crown scaling factor that does not correct for the number of trees sampled and (2) the persistent underestimate of the true VA due to a weak relationship between tree size and VA. The magnitude of differences between true densities and MHVD estimates suggest caution in using results based on the MHVD to inform management and restoration practices in the conifer forests of the American West.