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Thomas T. Veblen

Publications and source records attributed to Thomas T. Veblen.

15 recordsLinked to original sources

Linking seed size and number to trait syndromes in trees

Aim Our understanding of the mechanisms that maintain forest diversity under changing climate can benefit from knowledge about traits that are closely linked to fitness. We tested whether the link between traits and seed number and seed size is consistent with two hypotheses, termed the leaf economics spectrum and the plant size syndrome, or whether reproduction represents an independent dimension related to a seed size–seed number trade-off. Location Most of the data come from Europe, North and Central America and East Asia. A minority of the data come from South America, Africa and Australia. Time period 1960–2022. Major taxa studied Trees. Methods We gathered 12 million observations of the number of seeds produced in 784 tree species. We estimated the number of seeds produced by individual trees and scaled it up to the species level. Next, we used principal components analysis and generalized joint attribute modelling (GJAM) to map seed number and size on the tree traits spectrum. Results Incorporating seed size and number into trait analysis while controlling for environment and phylogeny with GJAM exposes relationships in trees that might otherwise remain hidden. Production of the large total biomass of seeds [product of seed number and seed size; hereafter, species seed productivity (SSP)] is associated with high leaf area, low foliar nitrogen, low specific leaf area (SLA) and dense wood. Production of high seed numbers is associated with small seeds produced by nutrient-demanding species with softwood, small leaves and high SLA. Trait covariation is consistent with opposing strategies: one fast-growing, early successional, with high dispersal, and the other slow-growing, stress-tolerant, that recruit in shaded conditions. Main conclusions Earth system models currently assume that reproductive allocation is indifferent among plant functional types. Easily measurable seed size is a strong predictor of the seed number and species seed productivity. The connection of SSP with the functional traits can form the first basis of improved fecundity prediction across global forests.

Global Ecology and Biogeography

Masting is shaped by tree-level attributes and stand structure, more than climate, in a Rocky Mountain conifer species

Masting describes the spatiotemporal variability in seed production by a population of plants. Both abiotic and biotic factors drive masting, but the importance of these factors can vary among individuals and populations. To better understand how a changing climate, altered disturbance regimes, or novel management strategies might affect future seed production, we quantified the joint influence of multiple factors on annual cone production in a widespread conifer species, Rocky Mountain ponderosa pine ( Pinus ponderosa var . scopulorum ). We reconstructed individual-level annual cone production across climatic gradients using the cone abscission scar method, and explored the causes and drivers of masting in this species. Site-level responses between weather and masting were highly variable, notably differing in the strength of their response to either summer or spring weather. In addition, the relationship to summer precipitation during cone initiation, a primary driver of annual seed production in this species, was strongest at hotter and drier sites. Additionally, we found that masting was strongly influenced by tree- and stand-level factors such as diameter, age, and local neighborhood density, all of which were associated with the mean, interannual variability, and between-tree synchrony of cone production at the individual-level. Larger and older trees produced more cones, more frequently, and with less synchrony than smaller and younger trees. Open grown trees experiencing lower levels of neighborhood competition also produced more cones with less interannual variability, but with higher between-tree synchrony. Because tree- and stand-level traits appear to regulate seed production more strongly than climate or weather in this species, management interventions targeting these factors could be powerful tools to manage future tree recruitment. Thus, current efforts to reduce stand density and conserve large trees in some ponderosa pine forests may enhance tree-level seed production and reduce variability in seed crops among years.

Arizona, Colorado, New Mexico, South Dakota, Wyomi

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

Nature Communications

Globally, tree fecundity exceeds productivity gradients

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Ecology Letters

North American tree migration paced by climate in the West, lagging in the East

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

PNAS

The effects of ENSO and the North American monsoon on mast seeding in two Rocky Mountain conifer species

We aimed to disentangle the patterns of synchronous and variable cone production (i.e. masting) and its relationship to climate in two conifer species native to dry forests of western North America. We used cone abscission scars to reconstruct ca 15 years of recent cone production in Pinus edulis and Pinus ponderosa , and used redundancy analysis to relate time series of annual cone production to climate indices describing the North American monsoon and the El Niño Southern Oscillation (ENSO). We show that the sensitivity to climate and resulting synchrony in cone production varies substantially between species. Cone production among populations of P. edulis was much more spatially synchronous and more closely related to large-scale modes of climate variability than among populations of P. ponderosa . Large-scale synchrony in P. edulis cone production was associated with the North American monsoon and we identified a dipole pattern of regional cone production associated with ENSO phase. In P. ponderosa , these climate indices were not strongly associated with cone production, resulting in asynchronous masting patterns among populations. This study helps frame our understanding of mast seeding as a life-history strategy and has implications for our ability to forecast mast years in these species.

Philosophical Transactions of the Royal Society B:

Is there tree senescence? The fecundity evidence

Despite its importance for forest regeneration, food webs, and human economies, changes in tree fecundity with tree size and age remain largely unknown. The allometric increase with tree diameter assumed in ecological models would substantially overestimate seed contributions from large trees if fecundity eventually declines with size. Current estimates are dominated by overrepresentation of small trees in regression models. We combined global fecundity data, including a substantial representation of large trees. We compared size–fecundity relationships against traditional allometric scaling with diameter and two models based on crown architecture. All allometric models fail to describe the declining rate of increase in fecundity with diameter found for 80% of 597 species in our analysis. The strong evidence of declining fecundity, beyond what can be explained by crown architectural change, is consistent with physiological decline. A downward revision of projected fecundity of large trees can improve the next generation of forest dynamic models.

Proceedings of the National Academy of Sciences

Continent-wide tree fecundity driven by indirect climate effects

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

Nature Communications

Climate, environment, and disturbance history govern resilience of western North American Forests

Before the advent of intensive forest management and fire suppression, western North American forests exhibited a naturally occurring resistance and resilience to wildfires and other disturbances. Resilience, which encompasses resistance, reflects the amount of disruption an ecosystem can withstand before its structure or organization qualitatively shift to a different basin of attraction. In fire-maintained forests, resilience to disturbance events arose primarily from vegetation pattern-disturbance process interactions at several levels of organization. Using evidence from 15 ecoregions, spanning forests from Canada to Mexico, we review the properties of forests that reinforced qualities of resilience and resistance. We show examples of multi-level landscape resilience, of feedbacks within and among levels, and how conditions have changed under climatic and management influences. We highlight geographic similarities and important differences in the structure and organization of historical landscapes, their forest types, and in the conditions that have changed resilience and resistance to abrupt or large-scale disruptions. We discuss the role of the regional climate in episodically or abruptly reorganizing plant and animal biogeography and forest resilience and resistance to disturbances. We give clear examples of these changes and suggest that managing for resilient forests is a construct that strongly depends on scale and human social values. It involves human communities actively working with the ecosystems they depend on, and the processes that shape them, to adapt landscapes, species, and human communities to climate change while maintaining core ecosystem processes and services. Finally, it compels us to embrace management approaches that incorporate ongoing disturbances and anticipated effects of climatic changes, and to support dynamically shifting patchworks of forest and non-forest. Doing so could make these shifting forest conditions and wildfire regimes less disruptive to individuals and society.

western North America

A statement of common ground regarding the role of wildfire in forested landscapes of the western United States

For millennia, wildfires have markedly influenced forests and non-forested landscapes of the western United States (US), and they are increasingly seen as having substantial impacts on society and nature. There is growing concern over what kinds and amounts of fire will achieve desirable outcomes and limit harmful effects on people and nature. Moreover, the increasing complexity surrounding cost and management of wildfires suggests that science should play a more prominent role in informing decisions about the need for fire in nature, and the need for society to adapt to the inevitable occurrence of different kinds and amounts of fire and smoke. Scientists widely view the natural wildfire regime as essential to western US forest ecosystem functioning. However, debates continue over how much low-, moderate-, and high severity fire is “natural” or desirable in these forests. Ongoing disagreement centers on the characteristics and importance of historical proportions and patch size distributions of low-, moderate-, and high-severity fires of dry, moist, and cold forests, and on the ecological consequences of changing fire-patch patterns and relative abundances. Scientists also debate the relative importance of climate and extreme weather versus fuel as drivers of high-severity fire, as well as the effectiveness and value of fuel treatments for reducing risks of undesired fire effects. Climate research shows that we should expect shifting future climates in all ecoregions. These expected changes make it difficult for scientists, land managers, and decision-makers to know the degree to which future forest management should be informed by historical conditions. There also is disagreement about how to make western forests more resilient to future disruptions in both climatic and fire regimes. To complicate matters, areas of scientific agreement -- the “common ground” shared by those in the research community -- are poorly articulated. Thus, the focus of the Fire Research Consensus (FRC) project has been to identify common ground among scientists, and provide a summary that can inform management. Land and fire managers are one audience for this report, as are stakeholders and the interested public. Our analysis, which results from extensive scientific literature reviews and questionnaires sent to western fire scientists and land managers, is summarized in nine key topics: A. Fire history and fire ecology vary with geography. B. Human impacts and management history vary with geography. C. Fire is a keystone process, which occurs in almost all western US forest types. D. Knowledge of historical range of variability (HRV) is useful but does not dictate land management goals. E. Forest structure, composition, and fuels have changed, affecting burn severity and fire extent. F. Climate and fuels both influence current fire sizes and their severities. G. The role of changing climatic conditions is increasingly important. H. Multiple fire ecology and fire history research approaches can be useful for characterizing fire regimes. I. Many existing fire management tools and strategies can be useful moving forward. We found much common ground that will be useful to scientists, managers, citizens, and policy decision-makers. For example, there is wide agreement among scientists that fire is one of the most essential influences on western forests and that more fire is needed on most landscapes, but not all wildfire behavior or extent will do. Fires can produce more positive benefits and fewer negative impacts when they burn with an ecologically appropriate mix and pattern of low, moderate, and high severity. Managers will need assistance and funding to create landscape conditions that favor more desirable fire behavior at broad spatial scales. Note that much societal impact from western wildfires occurs in non-forested landscapes that are not covered in this report, where findings would differ from those reported here for forested landscapes. We summarize additional key points below.

Western United States

Seed origin and warming constrain lodgepole pine recruitment, slowing the pace of population range shifts

Understanding how climate warming will affect the demographic rates of different ecotypes is critical to predicting shifts in species distributions. Here we present results from a common garden, climate change experiment in which we measured seedling recruitment of lodgepole pine, a widespread North American conifer that is also planted globally. Seeds from a low-elevation provenance had greater recruitment to their third year (by 323%) than seeds from a high-elevation provenance across sites within and above its native elevation range and across climate manipulations. Heating reduced (by 49%) recruitment to the third year of both low- and high-elevation seed sources across the elevation gradient, while watering alleviated some of the negative effects of heating (108% increase in watered plots). Demographic models based on recruitment data from the climate manipulations and long-term observations of adult populations revealed that heating could effectively halt modeled upslope range expansion except when combined with watering. Simulating fire and rapid post-fire forest recovery at lower elevations accelerated lodgepole pine expansion into the alpine, but did not alter final abundance rankings among climate scenarios. Regardless of climate scenario, greater recruitment of low-elevation seeds compensated for longer dispersal distances to treeline, assuming colonization was allowed to proceed over multiple centuries. Our results show that ecotypes from lower elevations within a species’ range could enhance recruitment and facilitate upslope range shifts with climate change.

Global Change Biology

Declines in low-elevation subalpine tree populations outpace growth in high-elevation populations with warming

Species distribution shifts in response to climate change require that recruitment increase beyond current range boundaries. For trees with long life spans, the importance of climate-sensitive seedling establishment to the pace of range shifts has not been demonstrated quantitatively. Using spatially explicit, stochastic population models combined with data from long-term forest surveys, we explored whether the climate-sensitivity of recruitment observed in climate manipulation experiments was sufficient to alter populations and elevation ranges of two widely distributed, high-elevation North American conifers. Empirically observed, warming-driven declines in recruitment led to rapid modelled population declines at the low-elevation, ‘warm edge’ of subalpine forest and slow emergence of populations beyond the high-elevation, ‘cool edge’. Because population declines in the forest occurred much faster than population emergence in the alpine, we observed range contraction for both species. For Engelmann spruce, this contraction was permanent over the modelled time horizon, even in the presence of increased moisture. For limber pine, lower sensitivity to warming may facilitate persistence at low elevations – especially in the presence of increased moisture – and rapid establishment above tree line, and, ultimately, expansion into the alpine. Synthesis . Assuming 21st century warming and no additional moisture, population dynamics in high-elevation forests led to transient range contractions for limber pine and potentially permanent range contractions for Engelmann spruce. Thus, limitations to seedling recruitment with warming can constrain the pace of subalpine tree range shifts.

Journal of Ecology

Biogeochemistry of beetle-killed forests: Explaining a weak nitrate response

A current pine beetle infestation has caused extensive mortality of lodgepole pine ( Pinus contorta ) in forests of Colorado and Wyoming; it is part of an unprecedented multispecies beetle outbreak extending from Mexico to Canada. In United States and European watersheds, where atmospheric deposition of inorganic N is moderate to low (<10 kg⋅ha⋅y), disturbance of forests by timber harvest or violent storms causes an increase in stream nitrate concentration that typically is close to 400% of predisturbance concentrations. In contrast, no significant increase in streamwater nitrate concentrations has occurred following extensive tree mortality caused by the mountain pine beetle in Colorado. A model of nitrate release from Colorado watersheds calibrated with field data indicates that stimulation of nitrate uptake by vegetation components unaffected by beetles accounts for significant nitrate retention in beetle-infested watersheds. The combination of low atmospheric N deposition (<10 kg⋅ha⋅y), tree mortality spread over multiple years, and high compensatory capacity associated with undisturbed residual vegetation and soils explains the ability of these beetle-infested watersheds to retain nitrate despite catastrophic mortality of the dominant canopy tree species.

Colorado

Climatic influences on the growth of subalpine trees in the Colorado Front Range

We examined variations in tree growth responses to climatic variations among different tree species and habitat types in the subalpine zone of the Colorado Front Range. We constructed 25 tree ring site chronologies (11 of Picea engelmannii , 9 of Abies lasiocarpa , 4 of Pinus contorta var. latifolia , and 1 of Pinus flexilis ) from a series of subalpine habitats ranging from xeric to wet. To establish tree growth responses to climatic variation, we used correlation and response function analyses to compare variations in ring widths with monthly temperature and precipitation records. At the driest sites, growth of Picea and Abies tracked climatic variation similarly. At mesic and wet sites, however, these species differed in their responses to climatic variation. The responses of Pinus contorta , sampled over a narrower range of habitat types, differed from those of Picea and Abies but did not differ among sites. Steep environmental gradients in the subalpine zone of the Front Range accounted for most of the observed differences in growth responses to climatic variation. Even at adjacent sites that differ only slightly in topographic position, tree growth responses to climatic variation were distinct. Interspecific differences in response to climatic variations generally were less important than site differences. Intersite differences in tree growth responses to climatic variation can be used as indicators of environmental differences among subalpine habitats.

Ecology

Disturbance regime and disturbance interactions in Rocky Mountain subalpine forest

1 The spatial and temporal patterns of fire, snow avalanches and spruce beetle out-breaks were investigated in Marvine Lakes Valley in the Colorado Rocky Mountains in forests of Picea engelmannii , Abies lasiocarpa , Pseudotsuga menziesii and Populus tremuloides . Dates and locations of disturbances were determined by dendrochronological techniques. A geographic information system (GIS) was used to calculate areas affected by the different disturbance agents and to examine the spatial relationships of the different disturbances. 2 In the Marvine Lakes Valley, major disturbance was caused by fire in the 1470s, the 1630s and the 1870s and by spruce beetle outbreak in c. 1716, 1827 and 1949. 3 Since c. 1633, 9% of the Marvine Lakes Valley has been affected by snow avalanches, 38.6% by spruce beetle outbreak and 59.1% by fire. At sites susceptible to avalanches, avalanches occur at a near-annual frequency. The mean return intervals for fire and spruce beetle outbreaks are 202 and 116.5 years, respectively. Turnover times for fire and spruce beetle outbreaks are 521 and 259 years, respectively. 4 Several types of disturbance interaction were identified. For example, large and severe snow avalanches influence the spread of fire. Similarly, following a stand-devastating fire or avalanche, Picea populations will not support a spruce beetle outbreak until individual trees reach a minimum diameter which represents at least 70 years' growth. Thus, recent fires and beetle outbreaks have nonoverlapping distributions.

Journal of Ecology