Explore the vast range of titles available.

The loss of large old trees in many ecosystems around the world poses a threat to ecosystem integrity. Large old trees are among the biggest organisms on Earth. They are keystone structures in forests, woodlands, savannas, agricultural landscapes, and urban areas, playing unique ecological roles not provided by younger, smaller trees. However, populations of large old trees are rapidly declining in many parts of the world, with serious implications for ecosystem integrity and biodiversity.

Ecological memory is central to how ecosystems respond to disturbance and is maintained by two types of legacies -information and material. Species life-history traits represent an adaptive response to disturbance and are an information legacy; in contrast, the abiotic and biotic structures (such as seeds or nutrients) produced by single disturbance events are material legacies. Disturbance characteristics that support or maintain these legacies enhance ecological resilience and maintain a \"safe operating space\" for ecosystem recovery. However, legacies can be lost or diminished as disturbance regimes and environmental conditions change, generating a \"resilience debt\" that manifests only after the system is disturbed. Strong effects of ecological memory on post-disturbance dynamics imply that contingencies (effects that cannot be predicted with certainty) of individual disturbances, interactions among disturbances, and climate variability combine to affect ecosystem resilience. We illustrate these concepts and introduce a novel ecosystem resilience framework with examples of forest disturbances, primarily from North America. Identifying legacies that support resilience in a particular ecosystem can help scientists and resource managers anticipate when disturbances may trigger abrupt shifts in forest ecosystems, and when forests are likely to be resilient.

The majority of the world's forests are used for multiple purposes, which often include the potentially conflicting goals of timber production and biodiversity conservation. A scientifically validated management approach that can reduce such conflicts is retention forestry, an approach modeled on natural processes, which emerged in the last 25 years as an alternative to clearcutting. A portion of the original stand is left unlogged to maintain the continuity of structural and compositional diversity. We detail retention forestry's ecological role, review its current practices, and summarize the large research base on the subject. Retention forestry is applicable to all forest biomes, complements conservation in reserves, and represents bottom-up conservation through forest manager involvement. A research challenge is to identify thresholds for retention amounts to achieve desired outcomes. We define key issues for future development and link retention forestry with land-zoning allocation at various scales, expanding its uses to forest restoration and the management of uneven—age forests.

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.

Persistent changes in tree mortality rates can alter forest structure, composition, and ecosystem services such as carbon sequestration. Our analyses of longitudinal data from unmanaged old forests in the western United States showed that background (noncatastrophic) mortality rates have increased rapidly in recent decades, with doubling periods ranging from 17 to 29 years among regions. Increases were also pervasive across elevations, tree sizes, dominant genera, and past fire histories. Forest density and basal area declined slightly, which suggests that increasing mortality was not caused by endogenous increases in competition. Because mortality increased in small trees, the overall increase in mortality rates cannot be attributed solely to aging of large trees. Regional warming and consequent increases in water deficits are likely contributors to the increases in tree mortality rates.

Large trees with cavities provide critical ecological functions in forests worldwide, including vital nesting and denning resources for many species. However, many ecosystems are experiencing increasingly rapid loss of large trees or a failure to recruit new large trees or both. We quantify this problem in a globally iconic ecosystem in southeastern Australia--forests dominated by the world's tallest angiosperms, Mountain Ash (Eucalyptus regnans). Tree, stand and landscape-level factors influencing the death and collapse of large living cavity trees and the decay and collapse of dead trees with cavities are documented using a suite of long-term datasets gathered between 1983 and 2011. The historical rate of tree mortality on unburned sites between 1997 and 2011 was >14% with a mortality spike in the driest period (2006-2009). Following a major wildfire in 2009, 79% of large living trees with cavities died and 57-100% of large dead trees were destroyed on burned sites. Repeated measurements between 1997 and 2011 revealed no recruitment of any new large trees with cavities on any of our unburned or burned sites. Transition probability matrices of large trees with cavities through increasingly decayed condition states projects a severe shortage of large trees with cavities by 2039 that will continue until at least 2067. This large cavity tree crisis in Mountain Ash forests is a product of: (1) the prolonged time required (>120 years) for initiation of cavities; and (2) repeated past wildfires and widespread logging operations. These latter factors have resulted in all landscapes being dominated by stands ≤72 years and just 1.16% of forest being unburned and unlogged. We discuss how the features that make Mountain Ash forests vulnerable to a decline in large tree abundance are shared with many forest types worldwide.

Variable retention harvesting evolved in the Douglas-fir region of the Pacific Northwest gradually in response to increasing dissatisfaction with the ecological consequences of clear-cutting, from the standpoint of wildlife habitat and other important forest functions. It is a harvesting technique that can provide for retention (continuity) of such structures as large and old live trees, snags, and logs. Variable retention is based on the natural model of the biological legacies that are typically left behind following natural disturbances, such as wildfire, wind, and flood. Variable retention is also an important technique for fulfilling the first silvicultural principle of ecological forestry, that of providing for continuity in structure, function, and composition between forest generations. The history and current application of variable retention approaches on forests in western Washington and Oregon states (USA), where many of the fundamental concepts were first developed and applied, is described in this article.

Quantifying historical fire regimes provides important information for managing contemporary forests. Historical fire frequency and severity can be estimated using several methods; each method has strengths and weaknesses and presents challenges for interpretation and verification. Recent efforts to quantify the timing of historical high-severity fire events in forests of western North America have assumed that the \"stand age\" variable from the US Forest Service Forest Inventory and Analysis (FIA) program reflects the timing of historical high-severity (i.e. stand-replacing) fire in ponderosa pine and mixed-conifer forests. To test this assumption, we re-analyze the dataset used in a previous analysis, and compare information from fire history records with information from co-located FIA plots. We demonstrate that 1) the FIA stand age variable does not reflect the large range of individual tree ages in the FIA plots: older trees comprised more than 10% of pre-stand age basal area in 58% of plots analyzed and more than 30% of pre-stand age basal area in 32% of plots, and 2) recruitment events are not necessarily related to high-severity fire occurrence. Because the FIA stand age variable is estimated from a sample of tree ages within the tree size class containing a plurality of canopy trees in the plot, it does not necessarily include the oldest trees, especially in uneven-aged stands. Thus, the FIA stand age variable does not indicate whether the trees in the predominant size class established in response to severe fire, or established during the absence of fire. FIA stand age was not designed to measure the time since a stand-replacing disturbance. Quantification of historical \"mixed-severity\" fire regimes must be explicit about the spatial scale of high-severity fire effects, which is not possible using FIA stand age data.

At global and regional scales, tree mortality rates are positively correlated with forest net primary productivity (NPP). Yet causes of the correlation are unknown, in spite of potentially profound implications for our understanding of environmental controls of forest structure and dynamics and, more generally, our understanding of broad-scale environmental controls of population dynamics and ecosystem processes. Here we seek to shed light on the causes of geographic patterns in tree mortality rates, and we consider some implications of the positive correlation between mortality rates and NPP. To reach these ends, we present seven hypotheses potentially explaining the correlation, develop an approach to help distinguish among the hypotheses, and apply the approach in a case study comparing a tropical and temperate forest. Based on our case study and literature synthesis, we conclude that no single mechanism controls geographic patterns of tree mortality rates. At least four different mechanisms may be at play, with the dominant mechanisms depending on whether the underlying productivity gradients are caused by climate or soil fertility. Two of the mechanisms are consequences of environmental selection for certain combinations of life-history traits, reflecting trade-offs between growth and defense (along edaphic productivity gradients) and between reproduction and persistence (as manifested in the adult tree stature continuum along climatic and edaphic gradients). The remaining two mechanisms are consequences of environmental influences on the nature and strength of ecological interactions: competition (along edaphic gradients) and pressure from plant enemies (along climatic gradients). For only one of these four mechanisms, competition, can high mortality rates be considered to be a relatively direct consequence of high NPP. The remaining mechanisms force us to adopt a different view of causality, in which tree growth rates and probability of mortality can vary with at least a degree of independence along productivity gradients. In many cases, rather than being a direct cause of high mortality rates, NPP may remain high in spite of high mortality rates. The independent influence of plant enemies and other factors helps explain why forest biomass can show little correlation, or even negative correlation, with forest NPP.