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USGS2021

The roles of environmental variation and parasite survival in virulence–transmission relationships

Disease outbreaks are a consequence of interactions among the three components of a host–parasite system: the infectious agent, the host and the environment. While virulence and transmission are widely investigated, most studies of parasite life-history trade-offs are conducted with theoretical models or tractable experimental systems where transmission is standardized and the environment controlled. Yet, biotic and abiotic environmental factors can strongly affect disease dynamics, and ultimately, host–parasite coevolution. Here, we review research on how environmental context alters virulence–transmission relationships, focusing on the off-host portion of the parasite life cycle, and how variation in parasite survival affects the evolution of virulence and transmission. We review three inter-related ‘approaches’ that have dominated the study of the evolution of virulence and transmission for different host–parasite systems: (i) evolutionary trade-off theory, (ii) parasite local adaptation and (iii) parasite phylodynamics. These approaches consider the role of the environment in virulence and transmission evolution from different angles, which entail different advantages and potential biases. We suggest improvements to how to investigate virulence–transmission relationships, through conceptual and methodological developments and taking environmental context into consideration. By combining developments in life-history evolution, phylogenetics, adaptive dynamics and comparative genomics, we can improve our understanding of virulence–transmission relationships across a diversity of host–parasite systems that have eluded experimental study of parasite life history.

Royal Society Open Science↗

USGS2022

An invasive prey provides long-lasting silver spoon effects for an endangered predator

The natal environment can have long-term fitness consequences for individuals, particularly via ‘silver spoon’ or ‘environmental matching’ effects. Invasive species could alter natal effects on native species by changing species interactions, but this potential remains unknown. Using 17 years of data on 2588 individuals across the entire US breeding range of the endangered snail kite ( Rostrhamus sociabilis ), a wetland raptor that feeds entirely on Pomacea snails, we tested for silver spoon and environmental matching effects on survival and movement and whether the invasion of a non-native snail may alter outcomes. We found support for silver spoon effects, not environmental matching, on survival that operated through body condition at fledging, explained by hydrology in the natal wetland. When non-native snails were present at the natal site, kites were in better condition, individual condition was less sensitive to hydrology, and kites fledged across a wider range of hydrologic conditions, leading to higher survival that persisted for at least 10 years. Movement between wetlands was driven by the current (adult) environment, and birds born in both invaded and uninvaded wetlands preferred to occupy invaded wetlands post-fledging. These results illustrate that species invasions may profoundly impact the role of natal environments on native species.

Florida↗

USGS1983

Clay mineral formation and transformation in rocks and soils

Three mechanisms for clay mineral formation (inheritance, neoformation, and transformation) operating in three geological environments (weathering, sedimentary, and diagenetic-hydrothermal) yield nine possibilities for the origin of clay minerals in nature. Several of these possibilities are discussed in terms of the rock cycle. The mineralogy of clays neoformed in the weathering environment is a function of solution chemistry, with the most dilute solutions favoring formation of the least soluble clays. After erosion and transportation, these clays may be deposited on the ocean floor in a lateral sequence that depends on floccule size. Clays undergo little reaction in the ocean, except for ion exchange and the neoformation of smectite; therefore, most clays found on the ocean floor are inherited from adjacent continents. Upon burial and heating, however, dioctahedral smectite reacts in the diagenetic environment to yield mixed-layer illite-smectite, and finally illite. With uplift and weathering, the cycle begins again. Refs.

Philosophical Transactions of the Royal Society A:↗

USGS1970

Nuclear methods applied to uranium geochemistry

Stable and radioactive daughter products produced from nuclear disintegrations of uranium have proved useful in fundamental studies of the geochemistry of uranium in igneous rock and sedimentary environments and in ore deposits. Information gained from geochemical studies of uranium migration has been used to develop models for attempts to date archeological, geological, and oceanographic environments represented by samples of bone, wood, charcoal, continental and marine carbonates, marine sediments, and glacially derived soils. Recent improvements of nuclear instrumentation and techniques allowed accurate measurements of natural radioactive isotopes, and it is now believed that radioactive equilibrium between the long-lived isotopes of the two uranium decay series is more the exception than the rule in nature. It was assumed that the 234U and 238U isotopes were in equilibrium until Thurber confirmed that considerable separation between 234U daughter and 238U parent exists in nature. It now has been documented that the 234U content may range from 60% deficient to 500% in excess relative to 238U. An excess of 15% of 234U isotope in sea water is well documented. A summary of previous work has shown that geochemical fractionation of the radioactive nuclides in 238U and 235U decay series takes place in the hydrologic environment, resulting in depletion of 230Th and 231Pa with respect to their parents, 238U, 234U, and 235U, in water and a complementary enrichment of these daughter nuclides in some sediments. Subsequent assimilation of Uranium, essentially free of radioactive daughters, occurs in some specific types of deposits such as carbonates and phosphates.

IEEE Transactions on Nuclear Science↗

USGS2020

Low stand density moderates growth declines during hot droughts in semi-arid forests

Increasing heat and aridity in coming decades is expected to negatively impact tree growth and threaten forest sustainability in dry areas. Maintaining low stand density has the potential to mitigate the negative effects of increasingly severe droughts by minimizing competitive intensity. However, the direct impact of stand density on the growing environment (i.e. soil moisture), and the specific drought metrics that best quantify that environment, are not well explored for any forest ecosystem. We examined the relationship of varying stand density (i.e. basal area) on soil moisture and stand‐level growth in a long‐term (multi‐decadal), ponderosa pine Pinus ponderosa , forest management experiment. We accounted for the influence of stand‐level density on moisture availability by measuring and modelling soil moisture using an ecosystem water balance model. To quantify the growing environment, we developed metrics of ecological drought that integrate the influence of moisture availability in the soil with moisture demand by the atmosphere. We paired these results with stand‐level dendrochronological data, avoiding the potential bias introduced from individual tree‐based assessments, and used critical climate period analysis to identify the timing and duration of these drought metrics that most relate to forest growth. We found that stand‐level growth is highly responsive to the combination of high temperature and low soil moisture. Growth in all stands was negatively related to temperature and positively related to moisture availability, although the sensitivity of growth to those conditions varied among stand density treatments. Growth enhancement during cool years is greatest in low density stands. In addition, low density stands displayed substantially higher long‐term average growth than higher density stands and maintained higher growth even when temperatures were high. Growth in low density stands also increased more than higher density stands in response to greater long‐term moisture availability. Synthesis and applications . We quantified the influence of stand‐level density on the environmental conditions that determine tree growth and related forest growth to patterns of moisture supply and demand. Our drought metrics, and analytical approach for quantifying drought impacts on forest growth, are a novel approach for assessing forest vulnerability to drought under climate change. These results provide new perspective on the potential for density management to mitigate drought stress and maintain forest stand growth during and after drought events in water‐limited forests.

Arizona↗

USGS2022

Methanogenic archaea in subsurface coal seams are biogeographically distinct: An analysis of metagenomically-derived mcrA sequences

The production of methane as an end-product of organic matter degradation in the absence of other terminal electron acceptors is common, and has often been studied in environments such as animal guts, soils, and wetlands due to its potency as a greenhouse gas. To date however, the study of the biogeographic distribution of methanogens across coal seam environments has been minimal. Here, we show that coal seams are host to a diverse range of methanogens, which are distinctive to each geological basin. Based on comparisons to close relatives from other methanogenic environments, the dominant methanogenic pathway in these basins is hydrogenotrophic, with acetoclastic being a second major pathway in the Surat Basin. Finally, mcrA and 16S rRNA gene primer biases were predominantly seen to affect the detection of Methanocellales, Methanomicrobiales and Methanosarcinales taxa in this study. Subsurface coal methanogenic community distributions and pathways presented here provide insights into important metabolites and bacterial partners for in situ coal biodegradation.

Environmental Microbiology↗

USGS2024

Common-garden experiment reveals outbreeding depression and region-of-origin effects on reproductive success in a frequently translocated tortoise

Human-mediated animal movement can expose wildlife populations to novel environments. Phenotypic plasticity can buffer against the challenges presented by novel environments, while adaptation to local ecosystems may limit resilience in novel ecosystems. Outbreeding depression during the mixing of disparate gene pools can also reduce reproductive success after long-distance movement. Here, we use a ‘common-garden’ population of gopher tortoises ( Gopherus polyphemus ), translocated from numerous sites across the state of Florida, USA, to a mitigation site in the north-west (panhandle) region to assess whether geographic origin, outbreeding effects, and behavioral plasticity influence reproductive success in this threatened keystone species. We found that females from north-east Florida produced clutches with lower hatching success than females from other regions. We detected regional differentiation in nest site selection behavior in the common environment of the translocation site, though these differences did not mediate the regional effect on hatching success. We also found evidence for outbreeding depression: hatching success declined with increasing parental geographic and genetic distances, dropping from 93% to 67% across the range of observed parental genetic distances. Together, these results suggest that newly admixed populations may suffer reproductive costs due to historical population differentiation, and that undetected outbreeding depression could significantly hamper conservation efforts for this species and others undergoing a variety of human-mediated movements.

Florida↗

USGS2014

What do we gain from simplicity versus complexity in species distribution models?

Species distribution models (SDMs) are widely used to explain and predict species ranges and environmental niches. They are most commonly constructed by inferring species' occurrence–environment relationships using statistical and machine-learning methods. The variety of methods that can be used to construct SDMs (e.g. generalized linear/additive models, tree-based models, maximum entropy, etc.), and the variety of ways that such models can be implemented, permits substantial flexibility in SDM complexity. Building models with an appropriate amount of complexity for the study objectives is critical for robust inference. We characterize complexity as the shape of the inferred occurrence–environment relationships and the number of parameters used to describe them, and search for insights into whether additional complexity is informative or superfluous. By building ‘under fit’ models, having insufficient flexibility to describe observed occurrence–environment relationships, we risk misunderstanding the factors shaping species distributions. By building ‘over fit’ models, with excessive flexibility, we risk inadvertently ascribing pattern to noise or building opaque models. However, model selection can be challenging, especially when comparing models constructed under different modeling approaches. Here we argue for a more pragmatic approach: researchers should constrain the complexity of their models based on study objective, attributes of the data, and an understanding of how these interact with the underlying biological processes. We discuss guidelines for balancing under fitting with over fitting and consequently how complexity affects decisions made during model building. Although some generalities are possible, our discussion reflects differences in opinions that favor simpler versus more complex models. We conclude that combining insights from both simple and complex SDM building approaches best advances our knowledge of current and future species ranges.

Ecography↗

USGS2021

New geochemical tools for investigating resource and energy functions at deep-sea cold seeps using amino-acid δ15N in chemosymbiotic mussels (Bathymodiolus childressi)

In order to reconstruct the ecosystem structure of chemosynthetic environments in the fossil record, geochemical proxies must be developed. Here, we present a suite of novel compound-specific isotope parameters for tracing chemosynthetic production with a focus on understanding nitrogen dynamics in deep-sea cold seep environments. We examined the chemosymbiotic bivalve Bathymodiolus childressi from three geographically distinct seep sites on the NE Atlantic Margin and compared isotope data to non-chemosynthetic littoral mussels to test whether water depth, seep activity, and/or mussel bed size are linked to differences in chemosynthetic production. The bulk isotope analysis of carbon (δ 13 C) and nitrogen (δ 15 N), and δ 15 N values of individual amino acids (δ 15 N AA ) in both gill and muscle tissues, as well as δ 15 N AA- derived parameters including trophic level (TL), baseline δ 15 N value (δ 15 N Phe ), and a microbial resynthesis index (Σ V ), were used to investigate specific geochemical signatures of chemosynthesis. Our results show that δ 15 N AA values provide a number of new proxies for relative reliance on chemosynthesis, including TL, ∑V, and both δ 15 N values and molar percentages (Gly/Glu mol% index) of specific AA. Together, these parameters suggested that relative chemoautotrophy is linked to both degree of venting from seeps and mussel bed size. Finally, we tested a Bayesian mixing model using diagnostic AA δ 15 N values, showing that percent contribution of chemoautotrophic versus heterotrophic production to seep mussel nutrition can be directly estimated from δ 15 N AA values. Our results demonstrate that δ 15 N AA analysis can provide a new set of geochemical tools to better understand mixotrophic ecosystem function and energetics, and suggest extension to the study of ancient chemosynthetic environments in the fossil record.

Geobiology↗

USGS2016

Additive impacts of experimental climate change increase risk to an ectotherm at the Arctic's edge

Globally, Arctic and Subarctic regions have experienced the greatest temperature increases during the last 30 years. These extreme changes have amplified threats to the freshwater ecosystems that dominate the landscape in many areas by altering water budgets. Several studies in temperate environments have examined the adaptive capacity of organisms to enhance our understanding of the potential repercussions of warming and associated accelerated drying for freshwater ecosystems. However, few experiments have examined these impacts in Arctic or Subarctic freshwater ecosystems, where the climate is changing most rapidly. To evaluate the capacity of a widespread ectotherm to anticipated environmental changes, we conducted a mesocosm experiment with wood frogs ( Rana sylvatica ) in the Canadian Subarctic. Three warming treatments were fully crossed with three drying treatments to simulate a range of predicted changes in wetland environments. We predicted wetland warming and drying would act synergistically, with water temperature partially compensating for some of the negative effects of accelerated drying. Across all drying regimes, a 1 °C increase in water temperature increased the odds of survival by 1.79, and tadpoles in 52-day and 64-day hydroperiod mesocosms were 4.1–4.3 times more likely to survive to metamorphosis than tadpoles in 45-day mesocosms. For individuals who survived to metamorphosis, there was only a weak negative effect of temperature on size. As expected, increased temperatures accelerated tadpole growth through day 30 of the experiment. Our results reveal that one of the dominant herbivores in Subarctic wetlands, wood frog tadpoles, are capable of increasing their developmental rates in response to increased temperature and accelerated drying, but only in an additive manner. The strong negative effects of drying on survival, combined with lack of compensation between these two environmental drivers, suggest changes in the aquatic environment that are expected in this ecosystem will reduce mean fitness of populations across the landscape.

Global Change Biology↗

USGS2016

Density-driven free-convection model for isotopically fractionated geogenic nitrate in sabkha brine

Subsurface brines with high nitrate (NO 3 − ) concentration are common in desert environments as atmospheric nitrogen is concentrated by the evaporation of precipitation and little nitrogen uptake. However, in addition to having an elevated mean concentration of ∼525 mg/L (as N), NO 3 − in the coastal sabkhas of Abu Dhabi is enriched in 15 N (mean δ 15 N ∼17‰), which is an enigma. A NO 3 − solute mass balance analysis of the sabkha aquifer system suggests that more than 90% of the nitrogen is from local atmospheric deposition and the remainder from ascending brine. In contrast, isotopic mass balances based on Δ 17 O, δ 15 N, and δ 18 O data suggest approximately 80 to 90% of the NO 3 − could be from ascending brine. As the sabkha has essentially no soil, no vegetation, and no anthropogenic land or water use, we propose to resolve this apparent contradiction with a density-driven free-convection transport model. In this conceptual model, the density of rain is increased by solution of surface salts, transporting near-surface oxygenated NO 3 − bearing water downward where it encounters reducing conditions and mixes with oxygen-free ascending geologic brines. In this environment, NO 3 − is partially reduced to nitrogen gas (N 2 ), thus enriching the remaining NO 3 − in heavy isotopes. The isotopically fractionated NO 3 − and nitrogen gas return to the near-surface oxidizing environment on the upward displacement leg of the free-convection cycle, where the nitrogen gas is released to the atmosphere and new NO 3 − is added to the system from atmospheric deposition. This recharge/recycling process has operated over many cycles in the 8000-year history of the shallow aquifer, progressively concentrating and isotopically fractionating the NO 3 − .

Groundwater↗

USGS2019

Characterization of the exoskeleton of the Antarctic king crab Paralomis birsteini

Ocean acidification is projected to inhibit the biogenic production of calcium-carbonate skeletons in marine organisms. Antarctic waters represent a natural environment in which to examine the long-term effects of carbonate undersaturation on calcification in marine predators. King crabs (Decapoda: Anomura: Lithodidae), which currently inhabit the undersaturated environment of the continental slope off Antarctica, are potential invasives on the Antarctic shelf as oceanic temperatures rise. Here, we describe the chemical, physical, and mechanical properties of the exoskeleton of the deep-water Antarctic lithodid Paralomis birsteini and compare our measurements with two decapod species from shallow water at lower latitudes: Cancer borealis (Brachyura: Cancridae) and Callinectes sapidus (Brachyura: Portunidae). Paralomis birsteini deposit proportionally more calcium carbonate in their predatory chelae than their protective carapaces, compared with the other two crab species. When exoskeleton thickness and microhardness were compared between the chelae and carapace, the magnitude of the difference between these body regions was significantly greater in P. birsteini than in the other species tested. Hence, there appeared to be a greater disparity in P. birsteini in overall investment in calcium-carbonate structures among regions of the exoskeleton. The imperatives of prey consumption and predator avoidance may be influencing the deposition of calcium to different parts of the exoskeleton in lithodids living in an environment undersaturated with respect to calcium carbonate.

Invertebrate Biology↗

USGS2011

Evaluating the demographic buffering hypothesis with vital rates estimated for Weddell seals from 30years of mark-recapture data

1. Life‐history theory predicts that those vital rates that make larger contributions to population growth rate ought to be more strongly buffered against environmental variability than are those that are less important. Despite the importance of the theory for predicting demographic responses to changes in the environment, it is not yet known how pervasive demographic buffering is in animal populations because the validity of most existing studies has been called into question because of methodological deficiencies. 2. We tested for demographic buffering in the southern‐most breeding mammal population in the world using data collected from 5558 known‐age female Weddell seals over 30 years. We first estimated all vital rates simultaneously with mark–recapture analysis and then estimated process variance and covariance in those rates using a hierarchical Bayesian approach. We next calculated the population growth rate’s sensitivity to changes in each of the vital rates and tested for evidence of demographic buffering by comparing properly scaled values of sensitivity and process variance in vital rates. 3. We found evidence of positive process covariance between vital rates, which indicates that all vital rates are affected in the same direction by changes in annual environment. Despite the positive correlations, we found strong evidence that demographic buffering occurred through reductions in variation in the vital rates to which population growth rate was most sensitive. Process variation in vital rates was inversely related to sensitivity measures such that variation was greatest in breeding probabilities, intermediate for survival rates of young animals and lowest for survival rates of older animals. 4. Our work contributes to a small but growing set of studies that have used rigorous methods on long‐term, detailed data to investigate demographic responses to environmental variation. The information from these studies improves our understanding of life‐history evolution in stochastic environments and provides useful information for predicting population responses to future environmental change. Our results for an Antarctic apex predator also provide useful baselines from a marine ecosystem when its top‐ and middle‐trophic levels were not substantially impacted by human activity.

Journal of Animal Ecology↗

USGS2012

ArxA, a new clade of arsenite oxidase within the DMSO reductase family of molybdenum oxidoreductases

Arsenotrophy, growth coupled to autotrophic arsenite oxidation or arsenate respiratory reduction, occurs only in the prokaryotic domain of life. The enzymes responsible for arsenotrophy belong to distinct clades within the DMSO reductase family of molybdenum-containing oxidoreductases: specifically arsenate respiratory reductase, ArrA, and arsenite oxidase, AioA (formerly referred to as AroA and AoxB). A new arsenite oxidase clade, ArxA, represented by the haloalkaliphilic bacterium Alkalilimnicola ehrlichii strain MLHE-1 was also identified in the photosynthetic purple sulfur bacterium Ectothiorhodospira sp. strain PHS-1. A draft genome sequence of PHS-1 was completed and an arx operon similar to MLHE-1 was identified. Gene expression studies showed that arx A was strongly induced with arsenite. Microbial ecology investigation led to the identification of additional arx A-like sequences in Mono Lake and Hot Creek sediments, both arsenic-rich environments in California. Phylogenetic analyses placed these sequences as distinct members of the ArxA clade of arsenite oxidases. ArxA-like sequences were also identified in metagenome sequences of several alkaline microbial mat environments of Yellowstone National Park hot springs. These results suggest that ArxA-type arsenite oxidases appear to be widely distributed in the environment presenting an opportunity for further investigations of the contribution of Arx-dependent arsenotrophy to the arsenic biogeochemical cycle.

Environmental Microbiology↗

USGS2006

Environmental framework of ground‐water contamination

Ramifications of contamination are increasingly involved in the majority of ground‐water problems. The volume of usable ground water is shrinking in many places because of dispersion of contaminated water . Consideration of ground‐water contamination as a multitude of independent problems, separately solvable as each problem arises, is outmoded; wise policies, relating water supply to contamination potential, are needed to alleviate and to forestall problems. Methodology of managing contamination problems calls for appropriate classification of the hydrogeologic environment; these classifications include aspects of interdependent factors such as permeability, sorption, hydraulic gradient, position of water table relative to some base, and distance from source of contamination . Effective evaluations relate the dynamics of the hydrogeologic environment to contingencies involving contamination , as man changes his water ‐development and waste‐disposal practices. Ways of contamination and pertinent parts of the physical environment include: waste‐disposal practices (at or near land surface and in deep formations), artificial recharge (at land surface and in aquifers), accidents, and salt‐ water contamination of aquifers (shallow depth from salty surface water and at variable depths from subjacent salty aquifers). Evaluation of waste‐disposal problems calls for appreciation of two opposing tendencies–the tendency of wastes to move with ground water and the tendency to be attenuated near disposal sites by decay or inherent decrease in potency, by chemical and physical sorption, and by dilution through dispersion of ground water . Mixed wastes of differing attenuation habits represent special complex problems.

Groundwater↗

USGS2006

Arsenic in ground water of the Western United States

Natural occurrences of ground water with moderate (10 to 50 micrograms per liter) to high (greater than 50 micrograms per liter) concentrations of arsenic are common throughout much of the Western United States. High concentrations of arsenic are generally associated with one of four geochemical environments: (1) basin-fill deposits of alluvial-lacustrine origin, particularly in semiarid areas, (2) volcanic deposits, (3) geothermal systems, and (4) uranium and gold-mining areas. These findings are based on an extensive literature review, compilation of unpublished reports and data, and the review of data bases containing more than 7,000 analyses of ground-water samples for arsenic. In the first two environments, arsenic appears to be associated with sediments derived, in part, from volcanic rocks of intermediate to acidic composition. Dissolved arsenic concentrations in water from volcanic aquifers in the same regions, however, may be low (less than 10 micrograms per liter). Solid phases (minerals, amorphous solids, and sedimentary organic matter) that supply the dissolved arsenic have not been identified in most areas. Alluvial and lacustrine sedimentary deposits appear to be an important source of arsenic in volcanic areas (such as Lane County, Oregon) and in areas underlain by basin-fill deposits (such as Carson Desert in Nevada and the Tulare Lake basin in California). Mobilization of arsenic in sedimentary aquifers may be, in part, a result of changes in the geochemical environment due to agricultural irrigation. In the deeper subsurface, elevated arsenic concentrations are associated with compaction caused by groundwater withdrawals.

Groundwater↗

USGS2014

Effect of light on biodegradation of Estrone, 17β-estradiol, and 17α-ethinylestradiol in stream sediment

Biodegradation of [A-ring 14 C] Estrone (E1), 17β-estradiol (E2), and 17α-ethinylestradiol (EE2) to 14 CO 2 was investigated under light and dark conditions in microcosms containing epilithon or sediment collected from Boulder Creek, Colorado. Mineralization of the estrogen A-ring was observed in all sediment treatments, but not epilithon treatments. No difference in net mineralization between light and dark treatments was observed for 14 C-E2. Net mineralization of 14 C-E1 and 14 C-EE2 was enhanced in light treatments. Extents of 14 CO 2 accumulation and rates of mineralization were significantly greater for E2 than E1 under dark conditions, but were comparable under light conditions. These results indicate substantial differences in the uptake and metabolism of E1 and E2 in the environment and suggest biorecalcitrance of E1 relative to E2 in light-limited environments. The extent of 14 CO 2 accumulation and rate of mineralization for EE2 in dark treatments were less than half of that observed for E2 and generally lower than for E1, consistent with previous reports of EE2 biorecalcitrance. However, 14 CO 2 accumulation and rates of mineralization were comparable for EE2, E2, and E1 under light conditions. These results indicate photoactivation and/or phototransformation/photodegradation processes can substantially enhance heterotrophic biodegradation of estrogens in sunlit environments and may play an important role in estrogen transport and attenuation.

Colorado↗

USGS2013

Tree growth and competition in an old-growth Picea abies forest of boreal Sweden: influence of tree spatial patterning

Question: What factors best characterize tree competitive environments in this structurally diverse old-growth forest, and do these factors vary spatially within and among stands? Location: Old-growth Picea abies forest of boreal Sweden. Methods: Using long-term, mapped permanent plot data augmented with dendrochronological analyses, we evaluated the effect of neighbourhood competition on focal tree growth by means of standard competition indices, each modified to include various metrics of trees size, neighbour mortality weighting (for neighbours that died during the inventory period), and within-neighbourhood tree clustering. Candidate models were evaluated using mixed-model linear regression analyses, with mean basal area increment as the response variable. We then analysed stand-level spatial patterns of competition indices and growth rates (via kriging) to determine if the relationship between these patterns could further elucidate factors influencing tree growth. Results: Inter-tree competition clearly affected growth rates, with crown volume being the size metric most strongly influencing the neighbourhood competitive environment. Including neighbour tree mortality weightings in models only slightly improved descriptions of competitive interactions. Although the within-neighbourhood clustering index did not improve model predictions, competition intensity was influenced by the underlying stand-level tree spatial arrangement: stand-level clustering locally intensified competition and reduced tree growth, whereas in the absence of such clustering, inter-tree competition played a lesser role in constraining tree growth. Conclusions: Our findings demonstrate that competition continues to influence forest processes and structures in an old-growth system that has not experienced major disturbances for at least two centuries. The finding that the underlying tree spatial pattern influenced the competitive environment suggests caution in interpreting traditional tree competition studies, in which tree spatial patterning is typically not taken into account. Our findings highlight the importance of forest structure – particularly the spatial arrangement of trees – in regulating inter-tree competition and growth in structurally diverse forests, and they provide insight into the causes and consequences of heterogeneity in this old-growth system.

Vï¿½sterbotten County↗