Understanding the interplay between environmental factors contributing to treeline formation and how these factors influence different life stages remains a major research challenge. We used an afforestation experiment including 92 000 trees to investigate the spatial and temporal dynamics of tree mortality and growth at treeline in the Swiss Alps. Seedlings of three high-elevation conifer species (Larix decidua, Pinus mugo ssp. uncinata, and Pinus cembra) were systematically planted along an altitudinal gradient at and above the current treeline (2075 to 2230 m above sea level [a.s.l.]) in 1975 and closely monitored during the following 30 years. We used decision-tree models and generalized additive models to identify patterns in mortality and growth along gradients in elevation, snow duration, wind speed, and solar radiation, and to quantify interactions between the different variables. For all three species, snowmelt date was always the most important environmental factor influencing mortality, and elevation was always the most important factor for growth over the entire period studied. Individuals of all species survived at the highest point of the afforestation for more than 30 years, although mortality was greater above 2160 m a.s.l., 50-100 m above the current treeline. Optimal conditions for height growth differed from those for survival in all three species: early snowmelt (ca. day of year 125-140 [where day 1 is 1 January]) yielded lowest mortality rates, but relatively later snowmelt (ca. day 145-150) yielded highest growth rates. Although snowmelt and elevation were important throughout all life stages of the trees, the importance of radiation decreased over time and that of wind speed increased. Our findings provide experimental evidence that tree survival and height growth require different environmental conditions and that even small changes in the duration of snow cover, in addition to changes in temperature, can strongly impact tree survival and growth patterns at treeline. Further, our results show that the relative importance of different environmental variables for tree seedlings changes during the juvenile phase as they grow taller.
Biological nitrogen fixation (BNF) is the major nitrogen (N) input in many terrestrial ecosystems, yet we know little about the mechanisms and feedbacks that control this process in natural ecosystems. We here examine BNF in four taxonomically and ecologically different groups over the course of forest ecosystem development. At nine sites along the Franz Josef soil chronosequence (South Westland, New Zealand) that range in age from 7 to 120000 yr old, we quantified BNF from the symbiotic plant Coriaria arborea, cyanolichens (primarily Pseudocyphellaria spp.), bryophytes (many species), and heterotrophic bacteria in leaf litter. We specifically examined whether these groups could act as "nitrostats" at the ecosystem level, turning BNF on when N is scarce (early in primary succession) and off when N is plentiful (later in succession and retrogression). Coriaria was abundant and actively fixing (approximately 11 kg N x ha(-1) x yr(-1)) in the youngest and most N-poor site (7 yr old), consistent with nitrostat dynamics. Coriaria maintained high BNF rates independent of soil N availability, however, until it was excluded from the community after a single generation. We infer that Coriaria is an obligate N fixer and that the nitrostat feedback is mechanistically governed by species replacement at the community level, rather than down-regulation of BNF at the physiological scale. Biological nitrogen fixation inputs from lichens (means of 0-2 kg N x ha(-1) x yr(-1)), bryophytes (0.7-10 kg N x ha(-1) x yr(-1)), and litter (1-2 kg N x ha(-1) x yr(-1)) were driven primarily by changes in density, which peaked at intermediate-aged sites (and increased with soil N availability) for both lichens and bryophytes, and grew monotonically with soil age (but did not change with soil N) for litter. This non-nitrostatic link between soil N availability and lichen/bryophyte BNF likely stems from increased tree biomass in more fertile sites, which increases epiphytic moisture conditions and habitable surface area. This apparent positive feedback could produce N-rich conditions.
Context dependency is deemed to position the outcomes of species interactions along a continuum of mutualism to parasitism. Thus, it is imperative to understand which factors determine where a particular interspecific interaction falls along the continuum. Over the past 20 years research on the ectomycorrhizal symbiosis has resulted in sufficient independent studies to now generalize about the factors and mechanisms that affect host response to ectomycorrhizas. Using meta-analysis we quantitatively evaluated the role of biotic (partner identity and colonization levels of ectomycorrhizal fungi) and abiotic (phosphorus levels) factors in determining host biomass, height, and shoot:root responses to ectomycorrhizal associations. On average, seedlings across multiple host genera increased in total biomass when inoculated with ectomycorrhizal fungi regardless of the identity of the fungal associate; host genera differed in the magnitude of response for both total biomass and shoot:root ratio. Association with different fungal genera modified only host allocation of biomass to shoots and roots. Neither level of colonization on inoculated seedlings nor the level of contamination on control seedlings relative to colonization levels by target fungi on inoculated seedlings was important in explaining variation in effect sizes for any growth response. None of our proposed factors (identity of partners, colonization level, magnitude of contamination, or duration of association) explained variation in effect sizes for shoot height, although in general seedlings were taller when inoculated with ectomycorrhizal fungi. Phosphorus additions did not influence effect sizes. Although the general trend across studies was for a positive response of hosts to ectomycorrhizal inoculation, publication bias and methodological issues effectively reduce and distort the spectrum on which we evaluate host responses to ectomycorrhizal inoculation. Our results indicate that the variation in ectomycorrhizal fungi perceived by the host may be of a discrete (presence/absence of ectomycorrhizal fungi) rather than continuous nature (variation in identity or abundance of ectomycorrhizal fungi).
Introduced species inevitably experience novel selection pressures in their new environments as a result of changes in mutualist and antagonist relationships. While most previous work has examined how escape from specialist enemies has influenced herbivore or pathogen resistance of exotic species, post-introduction shifts in exotic dependence on mutualists have not been considered. In a common environment, we compared dependence on AM fungi of North American and European populations of Hypericum perforatum (St. John's Wort), a forb native to Europe. Introduced North American populations responded less to inoculation with AM fungi than did European populations. Root architecture was strongly correlated with mycorrhizal response, and introduced populations had finer root architecture than native populations. Finally, introduced populations exhibited decreased root and increased reproductive allocation relative to European populations, consistent with a transition to a weedier life history; however, biomass allocation patterns were uncorrelated with mycorrhizal response. These findings are the first demonstration of a genetically based reduction of mycorrhizal dependence and shift in root architecture in an introduced species.
Population limitation is a fundamental tenet of ecology, but the relative roles of exogenous and endogenous mechanisms remain unquantified for most species. Here we used multi-model inference (MMI), a form of model averaging, based on information theory (Akaike's Information Criterion) to evaluate the relative strength of evidence for density-dependent and density-independent population dynamical models in long-term abundance time series of 1198 species. We also compared the MMI results to more classic methods for detecting density dependence: Neyman-Pearson hypothesis-testing and best-model selection using the Bayesian Information Criterion or cross-validation. Using MMI on our large database, we show that density dependence is a pervasive feature of population dynamics (median MMI support for density dependence = 74.7-92.2%), and that this holds across widely different taxa. The weight of evidence for density dependence varied among species but increased consistently, with the number of generations monitored. Best-model selection methods yielded similar results to MMI (a density-dependent model was favored in 66.2-93.9% of species time series), while the hypothesis-testing methods detected density dependence less frequently (32.6-49.8%). There were no obvious differences in the prevalence of density dependence across major taxonomic groups under any of the statistical methods used. These results underscore the value of using multiple modes of analysis to quantify the relative empirical support for a set of working hypotheses that encompass a range of realistic population dynamical behaviors.
Biodiversity is proposed to be important for the rate of ecosystem functions. Most biodiversity-ecosystem function studies, however, consider only one response variable at a time, and even when multiple variables are examined they are analyzed separately. This means that a very important aspect of biodiversity is overlooked: the possibility for different species to carry out different functions at any one time. We propose a conceptual model to explore the effects of species loss on overall ecosystem functioning, where overall functioning is defined as the joint effect of many ecosystem functions. We show that, due to multifunctional complementarity among species, overall functioning is more susceptible to species loss than are single functions. Modeled relationships between species richness and overall ecosystem functioning using five empirical data sets on monocultures reflected the range of effects of species loss on multiple functions predicted by the model. Furthermore, an exploration of the correlations across functions and the degree of redundancy within functions revealed that multifunctional redundancy was generally lower than single-function redundancy in these empirical data sets. We suggest that by shifting the focus to the variety of functions maintained by a diversity of species, the full importance of biodiversity for the functioning of ecosystems can be uncovered. Our results are thus important for conservation and management of biota and ecosystem services.
Climate change, with both warmer spring temperatures and greater temperature fluctuations, has altered phenologies, possibly leading to greater risk of spring frost damage to temperate deciduous woody plants. Phenological observations of 20 woody species from 1993 to 2012 in Trelease Woods, Champaign County, Illinois, USA, were used to identify years with frost damage to vegetative and reproductive phases. Local temperature records were used in combination with the phenological observations to determine what combinations of the two were associated with damage. Finally, a long-term temperature record (1889-1992) was evaluated to determine if the frequency of frost damage has risen in recent decades. Frost < or = -1.7 degrees C occurred after bud-break in 14 of the 20 years of observation. Frost damage occurred in five years in the interior and in three additional years at only the forest edge. The degree of damage varied with species, life stage, tissue (vegetative or reproductive), and phenological phase. Common features associated with the occurrence of damage to interior plants were (1) a period of unusual warm temperatures in March, followed by (2) a frost event in April with a minimum temperature < or = -6.1 degrees C with (3) a period of 16-33 days between the extremes. In the long-term record, 10 of 124 years met these conditions, but the yearly probability of frost damage increased significantly, from 0.03 during 1889-1979 to 0.21 during 1980-2012. When the criteria were "softened" to < or = -1.7 degrees C in April and an interval of 16-37 days, 31 of 124 years met the conditions, and the yearly damage probability increased significantly to 0.19 for 1889-1979 and 0.42 for 1980-2012. In this forest, the combination of warming trends and temperature variability (extremes) associated with climate change is having ecologically important effects, making previously rare frost damage events more common.
Interactions among multiple causes of ecological perturbation, such as climate change and disturbance, can produce "ecological surprises." Here, we examine whether climate-fire-vegetation interactions can produce ecological changes that differ in direction from those expected from the effects of climate change alone. To do so, we focus on the "Big Woods" of central Minnesota, USA, which was shaped both by climate and fire. The deciduous Big Woods forest replaced regional woodlands and savannas after the severity of regional fire regimes declined at ca. AD 1300. A trend toward wet conditions has long been assumed to explain the forest expansion, but we show that water levels at two lakes within the region (Wolsfeld Lake and Bufflehead Pond) were low when open woodlands were transformed into the Big Woods. Water levels were high instead at ca. 2240-795 BC when regional fire regimes were most severe. Based on the correlation between water levels and fire-regime severity, we infer that prolonged or repeated droughts after ca. AD 1265 reduced the biomass and connectivity of fine fuels (grasses) within the woodlands. As a result, regional fire severity declined and allowed tree populations to expand. Tree-ring data from the region show a peak in the recruitment of key Big Woods tree species during the AD 1930s drought and suggest that low regional moisture balance need not have been a limiting factor for forest expansion. The regional history, thus, demonstrates the types of counterintuitive ecosystem changes that may arise as climate changes in the future.
The neutral theory for community structure and biodiversity is dependent on the assumption that species are equivalent to each other in all important ecological respects. We explore what this concept of equivalence means in ecological communities, how such species may arise evolutionarily, and how the possibility of ecological equivalents relates to previous ideas about niche differentiation. We also show that the co-occurrence of ecologically similar or equivalent species is not incompatible with niche theory as has been supposed, because niche relations can sometimes favor coexistence of similar species. We argue that both evolutionary and ecological processes operate to promote the introduction and to sustain the persistence of ecologically similar and in many cases nearly equivalent species embedded in highly structured food webs. Future work should focus on synthesizing niche and neutral perspectives rather than dichotomously debating whether neutral or niche models provide better explanations for community structure and biodiversity.
Sustained whole-lake additions of 13C-enriched dissolved inorganic carbon (DIC), intended to increase experimentally the delta13C of DIC in the epilimnion of a small lake with high dissolved organic carbon (DOC), were made during three seasonal periods (spring, summer, and autumn). Coupled with carbon and nitrogen stable isotope analysis of zooplankton and several of their putative food sources, these additions were used to investigate seasonal changes in the relative contributions of different food sources to zooplankton diet in the lake. Four main potential food sources were considered: phytoplankton, heterotrophic bacteria (HB), methanotrophic bacteria (MOB), and green sulfur bacteria (GSB). Because the number of potential food sources exceeded the number of isotopes analyzed, a computer program (IsoSource) was used to estimate the range of possible contributions of the various food sources. During all three periods the added inorganic 13C quickly increased the epilimnetic DIC delta13C by between 18 per thousand and 21 per thousand above the initial value of approximately -21 per thousand. This 13C enrichment of DIC was rapidly transmitted to the particulate organic matter (POM), which included photosynthetic phytoplankton. In spring and summer, delta13C of both adult and juvenile Daphnia increased by approximately 10 per thousand, indicating that Daphnia utilized autochthonous carbon. However, this 13C labeling of Daphnia was not so obvious during the autumn period, when their delta13C generally decreased. According to the IsoSource model outputs based on both delta13C and delta15N values, Daphnia utilized all four potential food source types during spring, summer, and autumn, but in different proportions. The possible contribution of phytoplankton to Daphnia diet was substantial (25-71%) in all seasons. The possible contributions of the bacterial food sources were more variable. The possible contribution of GSB was minor (0-20%) at all times and negligible in autumn. The possible contribution of HB was higher but very variable. Methanotrophic bacteria always made a significant contribution to Daphnia diet and were likely the single most important food source in autumn. Since both HB and MOB in this high-DOC lake probably depend largely on allochthonous organic carbon, our results highlight the seasonal variability in the potential importance of ecosystem subsidies in lake food webs.
The "evolution of increased competitive ability" (EICA) hypothesis predicts that exotic species will adapt to reduced herbivore pressure by losing costly defenses in favor of competitive ability. Previous studies often support the prediction that plants from exotic populations will be less well defended than plants from native populations. However, results are mixed with respect to the question of whether plants from exotic populations have become more competitive. In a common-garden experiment involving plants from two native and two exotic populations of 14 different invasive species, we tested whether exotic plants generally grow larger than conspecific native plants, and whether patterns of relative growth depend on the intensity of competition. We found a quite consistent pattern of larger exotic than native plants, but only in the absence of competition. These results suggest that invasive species may often evolve increased growth, and that increased growth may facilitate adaptation to noncompetitive environments.
Tree architecture, growth, and mortality change with increasing tree size and associated light conditions. To date, few studies have quantified how size-dependent changes in growth and mortality rates co-vary with architectural traits, and how such size-dependent changes differ across species and possible light capture strategies. We applied a hierarchical Bayesian model to quantify size-dependent changes in demographic rates and correlated demographic rates and architectural traits for 145 co-occurring Malaysian rain-forest tree species covering a wide range of tree sizes. Demographic rates were estimated using relative growth rate in stem diameter (RGR) and mortality rate as a function of stem diameter. Architectural traits examined were adult stature measured as the 95-percentile of the maximum stem diameter (upper diameter), wood density, and three tree architectural variables: tree height, foliage height, and crown width. Correlations between demographic rates and architectural traits were examined for stem diameters ranging from 1 to 47 cm. As a result, RGR and mortality varied significantly with increasing stem diameter across species. At smaller stem diameters, RGR was higher for tall trees with wide crowns, large upper diameter, and low wood density. Increased mortality was associated with low wood density at small diameters, and associated with small upper diameter and wide crowns over a wide range of stem diameters. Positive correlations between RGR and mortality were found over the whole range of stem diameters, but they were significant only at small stem diameters. Associations between architectural traits and demographic rates were strongest at small stem diameters. In the dark understory of tropical rain forests, the limiting amount of light is likely to make the interspecific difference in the effects of functional traits on demography more clear. Demographic performance is therefore tightly linked with architectural traits such as adult stature, wood density, and capacity for horizontal crown expansion. The enhancement of a demographic trade-off due to interspecific variation in functional traits in the understory helps to explain species coexistence in diverse rain forests.
Seed dispersal has a powerful influence on population dynamics, genetic structuring, evolutionary rates, and community ecology. Yet, patterns of seed dispersal are difficult to measure due to methodological shortcomings in tracking dispersed seeds from sources of interest. Here we introduce a new method to track seed dispersal: stable isotope enrichment. It consists of leaf-feeding plants with sprays of 15N-urea during the flowering stage such that seeds developed after applications are isotopically enriched. We conducted a greenhouse experiment with Solanum americanum and two field experiments with wild Capsicum annuum in southern Arizona, USA, to field-validate the method. First, we show that plants sprayed with 15N-urea reliably produce isotopically enriched progeny, and that delta 15N (i.e., the isotopic ratio) of seeds and seedlings is a linear function of the 15N-urea concentration sprayed on mothers. We demonstrate that three urea dosages can be used to distinctly enrich plants and unambiguously differentiate their offspring after seeds are dispersed by birds. We found that, with high urea dosages, the resulting delta 15N values in seedlings are 10(3) - 10(4) times higher than the delta 15N values of normal plants. This feature allows tracking not only where seeds arrive, but in locations where seeds germinate and recruit, because delta 15N enrichment is detectable in seedlings that have increased in mass by at least two orders of magnitude before fading to normal delta 15N values. Last, we tested a mixing model to analyze seed samples in bulk. We used the delta 15N values of batches (i.e., combined seedlings or seeds captured in seed traps) to estimate the number of enriched seeds coming from isotopically enriched plants in the field. We confirm that isotope enrichment, combined with batch-sampling, is a cheap, reliable, and user-friendly method for bulk-processing seeds and is thus excellent for the detection of rare dispersal events. This method could further the study of dispersal biology, including the elusive, but critically important, estimation of long-distance seed dispersal.
Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (<1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C: N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N·ha-1·yr -1 above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.
In detritus-based ecosystems, autochthonous primary production contributes very little to the detritus pool. Yet primary producers may still influence the functioning of these ecosystems through complex interactions with decomposers and detritivores. Recent studies have suggested that, in aquatic systems, small amounts of labile carbon (C) (e.g., producer exudates), could increase the mineralization of more recalcitrant organic-matter pools (e.g., leaf litter). This process, called priming effect, should be exacerbated under low-nutrient conditions and may alter the nature of interactions among microbial groups, from competition under low-nutrient conditions to indirect mutualism under high-nutrient conditions. Theoretical models further predict that primary producers may be competitively excluded when allochthonous C sources enter an ecosystem. In this study, the effects of a benthic diatom on aquatic hyphomycetes, bacteria, and leaf litter decomposition were investigated under two nutrient levels in a factorial microcosm experiment simulating detritus-based, headwater stream ecosystems. Contrary to theoretical expectations, diatoms and decomposers were able to coexist under both nutrient conditions. Under low-nutrient conditions, diatoms increased leaf litter decomposition rate by 20% compared to treatments where they were absent. No effect was observed under high-nutrient conditions. The increase in leaf litter mineralization rate induced a positive feedback on diatom densities. We attribute these results to the priming effect of labile C exudates from primary producers. The presence of diatoms in combination with fungal decomposers also promoted decomposer diversity and, under low-nutrient conditions, led to a significant decrease in leaf litter C:P ratio that could improve secondary production. Results from our microcosm experiment suggest new mechanisms by which primary producers may influence organic matter dynamics even in ecosystems where autochthonous primary production is low.
Our objective was to infer the climate drivers of regionally synchronous fire years in dry forests of the U.S. northern Rockies in Idaho and western Montana. During our analysis period (1650-1900), we reconstructed fires from 9245 fire scars on 576 trees (mostly ponderosa pine, Pinus ponderosa P. & C. Lawson) at 21 sites and compared them to existing tree-ring reconstructions of climate (temperature and the Palmer Drought Severity Index [PDSI]) and large-scale climate patterns that affect modern spring climate in this region (El Niño Southern Oscillation [ENSO] and the Pacific Decadal Oscillation [PDO]). We identified 32 regional-fire years as those with five or more sites with fire. Fires were remarkably widespread during such years, including one year (1748) in which fires were recorded at 10 sites across what are today seven national forests plus one site on state land. During regional-fire years, spring-summers were significantly warm and summers were significantly warm-dry whereas the opposite conditions prevailed during the 99 years when no fires were recorded at any of our sites (no-fire years). Climate in prior years was not significantly associated with regional- or no-fire years. Years when fire was recorded at only a few of our sites occurred under a broad range of climate conditions, highlighting the fact that the regional climate drivers of fire are most evident when fires are synchronized across a large area. No-fire years tended to occur during La Niña years, which tend to have anomalously deep snowpacks in this region. However, ENSO was not a significant driver of regional-fire years, consistent with the greater influence of La Niña than El Niño conditions on the spring climate of this region. PDO was not a significant driver of past fire, despite being a strong driver of modern spring climate and modern regional-fire years in the northern Rockies.
Distance decay is used to describe the (usually exponential) decay in ecological similarity of assemblages between two sites as a function of their distance apart along an environmental gradient. Exponential distance-decay curves are routinely fitted by calculating the ecological similarity between each pair of sites, and fitting a linear regression to the points on a scatter plot of log-similarity vs. distance (x-axis). However, pairs of sites where the assemblages have no species in common pose a problem, because the similarity is zero, and the log transformation cannot be applied. Common fixes to this problem (i.e., either removing or transforming the zero values) are shown to have undesirable consequences and to give widely disparate estimates. A new method is presented as a special case of a generalized dissimilarity model. It is fitted very quickly and easily using existing software, and it does not require removal or transformation of the zero similarity points. Its simplicity makes it convenient for use in conjunction with the resampling methods that are routinely employed to test hypotheses, to obtain standard errors of estimated parameters, or to compare distance-decay curves. A word of caution about standard application of the bootstrap is noted, and modified bootstrap and jackknife alternatives are demonstrated.
Nestedness is a common biogeographic pattern in which small communities form proper subsets of large communities. However, the detection of nestedness in binary presence-absence matrices will be affected by both the metric used to quantify nestedness and the reference null distribution. In this study, we assessed the statistical performance of eight nestedness metrics and six null model algorithms. The metrics and algorithms were tested against a benchmark set of 200 random matrices and 200 nested matrices that were created by passive sampling. Many algorithms that have been used in nestedness studies are vulnerable to type I errors (falsely rejecting a true null hypothesis). The best-performing algorithm maintains fixed row and fixed column totals, but it is conservative and may not always detect nestedness when it is present. Among the eight indices, the popular matrix temperature metric did not have good statistical properties. Instead, the Brualdi and Sanderson discrepancy index and Cutler's index of unexpected presences performed best. When used with the fixed-fixed algorithm, these indices provide a conservative test for nestedness. Although previous studies have revealed a high frequency of nestedness, a reanalysis of 288 empirical matrices suggests that the true frequency of nested matrices is between 10% and 40%.
Long-term studies can broaden our ecological understanding and are particularly important when examining contingent effects that involve changes to dominance by long-lived species. Such a change occurred during the last century in Southwestern (USA) ponderosa pine (Pinus ponderosa) forests. We used five livestock grazing exclosures established in 1912 to quantify vegetation structure in 1941 and 2004. Our objectives were to (1) assess the effects of historical livestock grazing on overstory structure and age distribution, (2) assess the effects of recent livestock grazing and overstory on understory vegetation, and (3) quantify and explain changes in understory vegetation between 1941 and 2004. In 1941, canopy cover of tree regeneration was significantly higher inside exclosures. In 2004, total tree canopy cover was twice as high, density was three times higher, trees were smaller, and total basal area was 40% higher inside exclosures. Understory species density, herbaceous plant density, and herbaceous cover were negatively correlated with overstory vegetation in both years. Most understory variables did not differ between grazing treatments in 1941 but were lower inside exclosures in 2004. Differences between grazing treatments disappeared once overstory effects were accounted for, indicating that they were due to the differential overstory response to historical livestock grazing practices. Between 1941 and 2004, species density declined by 34%, herbaceous plant density by 37%, shrub cover by 69%, total herbaceous cover by 59%, graminoid cover by 39%, and forb cover by 82%. However, these variables did not differ between grazing treatments or years once overstory effects were accounted for, indicating that the declines were driven by the increased dominance of the overstory during this period. Our results demonstrate that historical livestock grazing practices are an aspect of land-use history that can affect ecosystem development. Grazing history must be considered when extrapolating results from one site to another. In addition, the understory vegetation was more strongly controlled by the ponderosa pine overstory than by recent livestock grazing or by temporal dynamics, indicating that overstory effects must be accounted for when examining understory responses in this ecosystem.
The Canadian High Arctic has been warming for several decades. Over this period, tundra plant communities have been influenced by regional climate change, as well as other disturbances. At a site on Ellesmere Island, Nunavut, Canada, we measured biomass and composition changes in a heath community over 13 years using a point-intercept method in permanent plots (1995-2007) and over 27 years using a biomass harvest comparison (1981-2008). Results from both methods indicate that the community became more productive over time, suggesting that this ecosystem is currently in transition. Bryophyte and evergreen shrub abundances increased, while deciduous shrub, forb, graminoid, and lichen cover did not change. Species diversity also remained unchanged. Because of the greater evergreen shrub cover, canopy height increased. From 1995 to 2007, mean annual temperature and growing season length increased at the site. Maximum thaw depth increased, while soil water content did not change. We attribute the increased productivity of this community to regional warming over the past 30-50 years. This study provides the first plot-based evidence for the recent pan-Arctic increase in tundra productivity detected by satellite-based remote-sensing and repeat-photography studies. These types of ground-level observations are critical tools for detecting and projecting long-term community-level responses to warming.
Determining the importance of independent variables is of practical relevance to ecologists and managers concerned with allocating limited resources to the management of natural systems. Although techniques that identify explanatory variables having the largest influence on the response variable are needed to design management actions effectively, the use of various indices to evaluate variable importance is poorly understood. Using Monte Carlo simulations, we compared six different indices commonly used to evaluate variable importance; zero-order correlations, partial correlations, semipartial correlations, standardized regression coefficients, Akaike weights, and independent effects. We simulated four scenarios to evaluate the indices under progressively more complex circumstances that included correlation between explanatory variables, as well as a spurious variable that was correlated with other explanatory variables, but not with the dependent variable. No index performed perfectly under all circumstances, but partial correlations and Akaike weights performed poorly in all cases. Zero-order correlations was the only measure that detected the presence of a spurious variable, whereas only independent effects assigned overlap areas correctly once the spurious variable was removed. We therefore recommend using zero-order correlations to eliminate predictor variables with correlations near zero, followed by the use of independent effects to assign overlap areas and rank variable importance.
We inferred climate drivers of 20th-century years with regionally synchronous forest fires in the U.S. northern Rockies. We derived annual fire extent from an existing fire atlas that includes 5038 fire polygons recorded from 12,070,086 ha, or 71% of the forested land in Idaho and Montana west of the Continental Divide. The 11 regional-fire years, those exceeding the 90th percentile in annual fire extent from 1900 to 2003 (>102,314 ha or approximately 1% of the fire atlas recording area), were concentrated early and late in the century (six from 1900 to 1934 and five from 1988 to 2003). During both periods, regional-fire years were ones when warm springs were followed by warm, dry summers and also when the Pacific Decadal Oscillation (PDO) was positive. Spring snowpack was likely reduced during warm springs and when PDO was positive, resulting in longer fire seasons. Regional-fire years did not vary with El Niño-Southern Oscillation (ENSO) or with climate in antecedent years. The long mid-20th century period lacking regional-fire years (1935-1987) had generally cool springs, generally negative PDO, and a lack of extremely dry summers; also, this was a period of active fire suppression. The climate drivers of regionally synchronous fire that we inferred are congruent with those of previous centuries in this region, suggesting a strong influence of spring and summer climate on fire activity throughout the 20th century despite major land-use change and fire suppression efforts. The relatively cool, moist climate during the mid-century gap in regional-fire years likely contributed to the success of fire suppression during that period. In every regional-fire year, fires burned across a range of vegetation types. Given our results and the projections for warmer springs and continued warm, dry summers, forests of the U.S. northern Rockies are likely to experience synchronous, large fires in the future.
Early statistical methods focused on pre-data probability statements (i.e., data as random variables) such as P values; these are not really inferences nor are P values evidential. Statistical science clung to these principles throughout much of the 20th century as a wide variety of methods were developed for special cases. Looking back, it is clear that the underlying paradigm (i.e., testing and P values) was weak. As Kuhn (1970) suggests, new paradigms have taken the place of earlier ones: this is a goal of good science. New methods have been developed and older methods extended and these allow proper measures of strength of evidence and multimodel inference. It is time to move forward with sound theory and practice for the difficult practical problems that lie ahead. Given data the useful foundation shifts to post-data probability statements such as model probabilities (Akaike weights) or related quantities such as odds ratios and likelihood intervals. These new methods allow formal inference from multiple models in the a prior set. These quantities are properly evidential. The past century was aimed at finding the "best" model and making inferences from it. The goal in the 21st century is to base inference on all the models weighted by their model probabilities (model averaging). Estimates of precision can include model selection uncertainty leading to variances conditional on the model set. The 21st century will be about the quantification of information, proper measures of evidence, and multi-model inference. Nelder (1999:261) concludes, "The most important task before us in developing statistical science is to demolish the P-value culture, which has taken root to a frightening extent in many areas of both pure and applied science and technology".
Ecological theory argues that the controls over ecosystem processes are structured hierarchically, with broader-scale drivers acting as constraints over the interactions and dynamics at nested levels of organization. In river ecosystems, these interactions may arise from broadscale variation in channel form that directly shapes benthic habitat structure and indirectly constrains resource supply and biological activity within individual reaches. To evaluate these interactions, we identified sediment characteristics, water chemistry, and denitrifier community structure as factors influencing benthic denitrification rates in a sixth-order river that flows through two physiographic provinces and the transitional zone between them, each with distinct geomorphological properties. We found that denitrification rates tracked spatial changes in sediment characteristics and varied seasonally with expected trends in stream primary production. Highest rates were observed during the spring and summer seasons in the physiographic province dominated by fine-grained sediments, illustrating how large-scale changes in river structure can constrain the location of denitrification hotspots. In addition, nirS and nirK community structure each responded differently to variation in channel form, possibly due to changes in dissolved oxygen and organic matter supply. This shift in denitrifier community structure coincident with higher rates of N removal via denitrification suggests that microbial community structure may influence biogeochemical processes.
Ectomycorrhizal fungi (EMF), a phylogenetically and physiologically diverse guild, form symbiotic associations with many trees and greatly enhance their uptake of nutrients and water. Elevated CO2, which increases plant carbon supply and demand for mineral nutrients, may change the composition of the EMF community, possibly altering nutrient uptake and ultimately forest productivity. To assess CO2 effects on EMF communities, we sampled mycorrhizae from the FACTS-I (Forest-Atmosphere Carbon Transfer and Storage) research site in Duke Forest, Orange County, North Carolina, USA, where Pinus taeda forest plots are maintained at either ambient or elevated CO2 (200 ppm above ambient) concentrations. Mycorrhizae were identified by DNA sequence similarity of the internal transcribed spacer ribosomal RNA gene region. EMF richness was very high; 72 distinct phylotypes were detected from 411 mycorrhizal samples. Overall EMF richness and diversity were not affected by elevated CO2, but increased CO2 concentrations altered the relative abundances of particular EMF taxa colonizing fine roots, increased prevalence of unique EMF species, and led to greater EMF community dissimilarity among individual study plots. Natural variation among plots in mean potential net nitrogen (N) mineralization rates was a key determinant of EMF community structure; increasing net N mineralization rate was negatively correlated with EMF richness and had differential effects on the abundance of particular EMF taxa. Our results predict that, at CO2 concentrations comparable to that predicted for the year 2050, EMF community composition and structure will change, but diversity will be maintained. In contrast, high soil N concentrations can negatively affect EMF diversity; this underscores the importance of considering CO2 effects on forest ecosystems in the context of background soil chemical parameters and other environmental perturbations such as acid deposition or fertilizer runoff.
Individual variance in lifetime fecundity within populations is a life-history parameter of crucial evolutionary and ecological significance. However, knowledge of its magnitude and underlying mechanisms in natural populations is biased toward short-lived taxa. This paper summarizes results of a 23-year study on a population of the Mediterranean shrub Lavandula latifolia. We document the within-population pattern of individual variation in instantaneous and lifetime fecundity (as estimated by inflorescence production) and explore the mechanisms producing the lognormal distribution of individual fecundities by means of an individual-based simulation model. Throughout the study period, a few individuals consistently produced most inflorescences while the majority of plants exhibited moderate-to-low fecundities. The shape of yearly distributions of annual fecundities varied little across years, and most annual fecundity distributions did not depart significantly from a lognormal. The distribution of individual lifetime fecundity did not depart from lognormality. Despite the simplicity of the premises of our simulation model, it was remarkably successful at predicting the shapes of fecundity distributions and the early establishment of a persistent fecundity hierarchy. The agreement between model results and empirical data supports the view that multiplicative interactions of randomly varying environmental effects can play a central role in determining individual variation in lifetime fecundity in L. latifolia, and suggests that environmental stochasticity can be decisive in the genesis of strong fecundity hierarchies in long-lived plants.
Climate models predict precipitation changes for much of the humid tropics, yet few studies have investigated the potential consequences of drought on soil carbon (C) cycling in this important biome. In wet tropical forests, drought could stimulate soil respiration via overall reductions in soil anoxia, but previous research suggests that litter decomposition is positively correlated with high rainfall fluxes that move large quantities of dissolved organic matter (DOM) from the litter layer to the soil surface. Thus, reduced rainfall could also limit C delivery to the soil surface, reducing respiration rates. We conducted a throughfall manipulation experiment to investigate how 25% and 50% reductions in rainfall altered both C movement into soils and the effects of those DOM fluxes on soil respiration rates. In response to the experimental drought, soil respiration rates increased in both the -25% and -50% treatments. Throughfall fluxes were reduced by 26% and 55% in the -25% and -50% treatments, respectively. However, total DOM fluxes leached from the litter did not vary between treatments, because the concentrations of leached DOM reaching the soil surface increased in response to the simulated drought. Annual DOM concentrations averaged 7.7 +/- 0.8, 11.2 +/- 0.9, and 15.8 +/- 1.2 mg C/L in the control, -25%, and -50% plots, respectively, and DOM concentrations were positively correlated with soil respiration rates. A laboratory incubation experiment confirmed the potential importance of DOM concentration on soil respiration rates, suggesting that this mechanism could contribute to the increase in CO2 fluxes observed in the reduced rainfall plots. Across all plots, the data suggested that soil CO2 fluxes were partially regulated by the magnitude and concentration of soluble C delivered to the soil, but also by soil moisture and soil oxygen availability. Together, our data suggest that declines in precipitation in tropical rain forests could drive higher CO2 fluxes to the atmosphere both via increased soil 02 availability and through responses to elevated DOM concentrations.
Environmental-stress-mediated geographic variation in reproductive parameters has been little studied in natural vertebrate populations outside the context of climatic variation. Based on life-history theory, an increase in the degree of environmental stress experienced by a population should lead to (1) a shift in reproductive allocation from fecundity to offspring quality, (2) stronger trade-offs between reproductive parameters, and (3) changes in the relationship between female phenotype and maternal investment. To test these predictions, we investigated geographic variation in maternal investment of moor frogs (Rana arvalis) in relation to breeding site acidity (pH 4-8). We found that mean egg size increased and clutch size and total reproductive output (TRO) decreased with increasing acidity among 19 Swedish moor frog populations. Tests for variation and co-variation in maternal investment and female size and age in 233 females from a subset of four acid origin (AO) and four neutral origin (NO) populations revealed that clutch size and TRO increased with female size in both acid and neutral environments. However, in AO populations, egg size also increased with female size, and clutch size and TRO with female age, whereas in NO populations, egg size increased with female age. The strength of the egg-size-clutch-size tradeoff tended to be stronger in AO than in NO females as expected if the former experience stronger environmental constraints. All in all, these results suggest that environmental acidification selects for investment in larger eggs at a cost to fecundity, imposes negative effects on reproductive output, and alters the relationship between female phenotype and maternal investment.
Low Ca/Mg ratios (a defining component of serpentine soils) and low water environmental conditions often co-occur in nature and are thought to exert strong selection pressures on natural populations. However, few studies test the individual and combined effects of these environmental factors. We investigated the effects of low Ca/Mg ratio and low water availability on plant leaf, stem, stolon, and floral traits of Mimulus guttatus, a bodenvag species, i.e., a species that occurs in serpentine and non-serpentine areas. We quantified genetic variation and genetic variation for plasticity for these leaf, stem, stolon, and floral traits at three hierarchical levels: field-habitat type, population, and family, and we evaluated the relative importance of local adaptation and plasticity. We chose two populations and 10 families per population from four distinct field "habitat types" in northern California: high Ca/Mg ratio (non-serpentine) and season-long water availability, high Ca/Mg ratio and seasonally drying, low Ca/Mg ratio (serpentine) and season-long water availability, and low Ca/Mg ratio and seasonally drying. Seedlings were planted into greenhouse treatments that mimicked the four field conditions. We only detected genetic variation for stem diameter and length of longest leaf at the field-habitat level, but we detected genetic variation at the family level for nearly all traits. Soil chemistry and water availability had strong phenotypic effects, alone and in combination. Our hypothesis of an association between responses to low water levels and low Ca/Mg ratio was upheld for length of longest leaf, stem diameter, corolla width, and total number of reproductive units, whereas for other traits, responses to Ca/Mg ratio and low water were clearly independent. Our results suggest that traits may evolve independently from Ca/Mg ratios and water availability and that our focal traits were not simple alternative measures of vigor. We found genetic variation for plasticity both at the field-habitat type and family levels for half of the traits studied. Phenotypic plasticity and genetic variation for plasticity appear to be more important than local adaptation in the success of these M. guttatus populations found across a heterogeneous landscape in northern California. Phenotypic plasticity is an important mechanism maintaining the broad ecological breadth of native populations of M. guttatus.
We report that iron-reducing bacteria are primary mediators of anaerobic carbon oxidation in upland tropical soils spanning a rainfall gradient (3500-5000 mm/yr) in northeast Puerto Rico. The abundant rainfall and high net primary productivity of these tropical forests provide optimal soil habitat for iron-reducing and iron-oxidizing bacteria. Spatially and temporally dynamic redox conditions make iron-transforming microbial communities central to the belowground carbon cycle in these wet tropical forests. The exceedingly high abundance of iron-reducing bacteria (up to 1.2 x 10(9) cells per gram soil) indicated that they possess extensive metabolic capacity to catalyze the reduction of iron minerals. In soils from the higher rainfall sites, measured rates of ferric iron reduction could account for up to 44% of organic carbon oxidation. Iron reducers appeared to compete with methanogens when labile carbon availability was limited. We found large numbers of bacteria that oxidize reduced iron at sites with high rates of iron reduction and large numbers of iron reducers. The coexistence of large populations of iron-reducing and iron-oxidizing bacteria is evidence for rapid iron cycling between its reduced and oxidized states and suggests that mutualistic interactions among these bacteria ultimately fuel organic carbon oxidation and inhibit CH4 production in these upland tropical forests.
Connectivity among populations and habitats is important for a wide range of ecological processes. Understanding, preserving, and restoring connectivity in complex landscapes requires connectivity models and metrics that are reliable, efficient, and process based. We introduce a new class of ecological connectivity models based in electrical circuit theory. Although they have been applied in other disciplines, circuit-theoretic connectivity models are new to ecology. They offer distinct advantages over common analytic connectivity models, including a theoretical basis in random walk theory and an ability to evaluate contributions of multiple dispersal pathways. Resistance, current, and voltage calculated across graphs or raster grids can be related to ecological processes (such as individual movement and gene flow) that occur across large population networks or landscapes. Efficient algorithms can quickly solve networks with millions of nodes, or landscapes with millions of raster cells. Here we review basic circuit theory, discuss relationships between circuit and random walk theories, and describe applications in ecology, evolution, and conservation. We provide examples of how circuit models can be used to predict movement patterns and fates of random walkers in complex landscapes and to identify important habitat patches and movement corridors for conservation planning.
The response of ecological communities to anthropogenic disturbance is of both scientific and practical interest. Communities where all species respond to disturbance in a similar fashion (synchrony) will exhibit large fluctuations in total biomass and dramatic changes in ecosystem function. Communities where some species increase in abundance while others decrease after disturbance (compensation) can maintain total biomass and ecosystem function in the face of anthropogenic change. We examined dynamics of the Little Rock Lake (Wisconsin, USA) zooplankton community in the context of an experimental pH manipulation conducted in one basin of the lake. A novel application of wavelets was used to partition patterns of synchrony and compensation by time scale. We find interestingly that some time series show both patterns of synchrony and compensation depending on the scale of analysis. Within the unmanipulated basin, we found subtle patterns of synchrony and compensation within the community, largely at a one-year time scale corresponding to seasonal variation. Within the acidified lake basin, dynamics shifted to longer time scales corresponding to the pattern of pH manipulation. Comparisons between pairs of species in different functional groups showed both strong compensatory and synchronous responses to disturbance. The strongest compensatory signal was observed for two species of Daphnia whose life history traits lead to synchrony at annual time scales, but whose differential sensitivity to acidification led to compensation at multiannual time scales. The separation of time scales inherent in the wavelet method greatly facilitated interpretation as patterns resulting from seasonal drivers could be separated from patterns driven by pH manipulation.
The link between biodiversity and ecosystem functioning is now well established, but the challenge remains to develop a mechanistic understanding of observed effects. Predator-prey interactions provide an opportunity to examine the role of resource partitioning, thought to be a principal mediator of biodiversity-function relationships. To date, interactions between multiple predators and their prey have typically been investigated in simplified agricultural systems with limited scope for resource partitioning. Thus there remains a dearth of studies examining the functional consequences of predator richness in diverse food webs. Here, we manipulated a species-rich intertidal food web, crossing predator diversity with total predator density, to simultaneously examine the independent and interactive effects of diversity and density on the efficiency of secondary resource capture. The effect of predator diversity was only detectable at high predator densities where competitive interactions between individual predators were magnified; the rate of resource capture within the species mixture more than doubled that of the best-performing single species. Direct observation of species-specific resource use in monoculture, as quantified by patterns of prey consumption, provided clear evidence that species occupied distinct functional niches, suggesting a mechanistic explanation of the observed diversity effect.
In this paper we quantify the rate of spread of the newly emerged pathogen Mycoplasma gallisepticum of the House Finch, Carpodacus mexicanus, in its introduced range. We compare and contrast the rapid, yet decelerating, rate of spread of the pathogen with the slower, yet accelerating rate of spread of the introduced host. Comparing the rate of spread of this pathogen to pathogens in terrestrial mammalian hosts, we see that elevation and factors relating to host abundance restrict disease spread, rather than finding any major effects of discrete barriers or anthropogenic movement. We examine the role of seasonality in the rate of spread, finding that the rate and direction of disease spread relates more to seasonality in host movement than to seasonality in disease prevalence. We conclude that asymptomatic carriers are major transmitters of Mycoplasma gallisepticum into novel locations, a finding which may also be true for many other diseases, such as West Nile Virus and avian influenza.
Iriartea deltoidea (Arecaceae) is an abundant canopy palm with a wide geographic distribution in Neotropical wet forests. We analyzed the genetic profile across three generations of Iriartea within a 43-ha area encompassing two areas of second-growth and adjoining old-growth forest at La Selva Biological Field Station in northeastern Costa Rica. A total of 311 reproductively mature trees, 99 large saplings, 207 small saplings, and 601 seedlings were genotyped using 141 AFLP loci. Parentage analysis revealed high dispersal distances, both for seed (over 2.3 km) and pollen (over 3.8 km), indicating a large genetic neighborhood within La Selva Biological Station. In a 20-ha area of second growth, the founding palm population was dominated by a small number of parental trees located in the adjacent old-growth forest; two old-growth trees contributed 48% of the second-growth genes. The genetic diversity of reproductively mature trees in this second-growth forest was significantly reduced compared to adjacent old-growth forest. Within 400 m of the border with old-growth forest, we observed a similar reduction of genetic diversity in saplings, and an even greater loss of genetic diversity in the second generation of seedlings. Nearly half of these seedlings were offspring of local parents. In contrast, in the distant portion of second-growth forest (400-800 m from the old-growth border), parentage analysis showed that 40% of seedlings originated from outside the study area and only 10% were offspring of local parents. These high levels of gene flow maintained genetic diversity in saplings and seedlings similar to levels observed in old-growth forest. Our findings highlight the importance of gene flow from diverse seed and pollen sources for sustaining levels of genetic diversity of tree populations in second-growth forests.
The scales of population structure in marine species depend on the degree to which larvae from different populations are mixed in the plankton. There is an intriguing trend in marine population genetic studies of significant genetic structure for larvae, recruits, or populations at fine scales that is unpatterned across space and changes through time. This "chaotic genetic patchiness" suggests that larval pools are not well mixed in the plankton. However, few studies have been able to distinguish among potential causes of spatial and temporal genetic heterogeneity: changes in larval migration patterns, changes in environmental selection, or stochasticity caused by "sweepstakes" reproductive success of spawners creating detectable family structure. Here we use microsatellite markers to show that significant allele frequency shifts occurred sporadically in space and time for cohorts of recruits of Paralabrax clathratus (kelp bass) collected once every two weeks over two years from five sites in the Santa Barbara Channel, California, USA. We found that the pattern of genetic differentiation among cohorts was explained by a combination of (1) family structure in some cohorts, evidenced by half and full siblings, and (2) an indication of changes in larval delivery. It is unlikely but possible that environmental selection also plays a role. Although sampling of potential source populations was incomplete, cohorts arriving during western current flows show most genetic similarity with a population sample collected in the west, and cohorts arriving during current flows from the southeast show similarity with population samples collected in the south and east. Despite the family structure apparent in some cohorts, these "sweepstakes" events occur on too fine a scale to create lasting year class genetic structure. The results corroborate oceanographic models of larval dispersal, which suggest that larval mixing in the plankton is less extensive than previously believed.
Competition-colonization trade-offs are theorized to be a mechanism of coexistence in communities structured by environmental fluctuations. But many studies that have tested for the trade-off have failed to detect it, likely because a spatiotemporally structured environment and many species assemblages are needed to adequately test for a competition-colonization trade-off. Here, we present a unique 32-year study of rock-dwelling lichens in New Mexico, USA, in which photographs were used to quantify lichen life history traits and interactions through time. These data allowed us to determine whether there were any trade-offs between traits associated with colonization and competition, as well as the relationship between diversity and disturbance in the community. We did not find evidence for a trade-off between competitive ability and colonization rate or any related life history traits. Interestingly, we did find a peak in all measures of species diversity at intermediate levels of disturbance, consistent with the intermediate disturbance hypothesis pattern. We suggest that the coexistence of the dominant species in this system is regulated by differences in persistence and growth rate mediating overgrowth competition rather than a competition-colonization trade-off.
Leaching is a mechanism for the release of nutrients from litter or senesced leaves that can drive interactions among plants, microbes, and soil. Although leaching is well established in conceptual models of litter decomposition, potential nutrient solubility of mineral elements from recently senesced litter has seldom been quantified. Using a standardized extraction (1:50 litter-to-water ratio and four-hour extraction) and recently senesced leaf litter of 41 tropical tree and liana species, we investigated how solubility varies among elements, and whether the solubility of elements could be predicted by litter traits (e.g., lignin, total element concentrations). In addition, we investigated nutrient forms (i.e., inorganic and organic) and ratios in leachate. Water-soluble elements per unit litter mass were strongly predicted by total initial litter element concentrations for potassium (K; r2 = 0.79), sodium (Na; r2 = 0.51) and phosphorus (P; r2 = 0.66), while a significant but weaker positive relationship was found for nitrogen (N; r2 = 0.36). There was no significant relationship for carbon (C) or calcium (Ca). Element-specific solubility varied markedly. On average 100% of total K, 35% of total P, 28% of total Na, 5% of total N, 4% of total Ca, and 3% of total C were soluble. For soluble P, 90% was inorganic orthophosphate. The high solubility of K, Na, and P as inorganic orthophosphate suggests that these nutrients can become rapidly available to litter microbes with no metabolic cost. Few common predictors of decomposition rates were correlated with element solubility, although soluble C (milligrams per gram of litter) was negatively related to lignin content (r2 = 0.19; P < 0.004). Solubility of elements was linked within a species: when a species ranked high in the soluble fraction of one element, it also ranked high in the solubility of other elements. Overall nutrient-specific patterns of solubility from recently senesced litter emphasize that litter elements cannot be treated equally in our conceptual and empirical models of decomposition. The relatively high potential solubility of P as orthophosphate from fresh litter advances our understanding of ecological stoichiometric ratios and nutrient bioavailability in tropical forests.
The pine-dominated forests of west-central Mexico are internationally recognized for their high biodiversity, and some areas are protected through various conservation measures including prohibition of human activity. In this region, however, there is evidence for human settlement dating back to ca. AD 1200. It is therefore unclear whether the present forest composition and structure are part of a successional stage following use by indigenous human populations during the past, or due to natural processes, such as climate. We present a study reconstructing the vegetation dynamics of pine-dominated forest over the past 4200 years using paleoecological techniques. Results from fossil pollen and charcoal indicate that, in this region, pine-dominated forests are the native vegetation type and not anthropogenically derived secondary succession. The predominant driving mechanism for the expansion of pine-dominated forest appears to be intervals of aridity and naturally induced burning. A close association is noted between pine abundance and longer-term climatic trends, including intervals of aridity between ca. 4200 and 2500, 1200 and 850, and 500 and 200 cal yr BP and shorter-term trends. Evident periodicity occurs in pine and Poaceae abundance every 80 years. These short-term quasi-periodic oscillations have been recorded in a number of lake and ocean sediments in Mexico and are thought to be linked to solar forcing resulting in drought cycles that occur at approximately the same time intervals.
A critical assumption underlying terrestrial ecosystem models is that soil microbial communities, when placed in a common environment, will function in an identical manner regardless of the composition of that community. Given high species diversity in microbial communities and the ability of microbes to adapt rapidly to new conditions, this assumption of functional redundancy seems plausible. We test the assumption by comparing litter decomposition rates in experimental microcosms inoculated with distinct microbial communities. We find that rates of carbon dioxide production from litter decomposition were dependent upon the microbial inoculum, with differences in the microbial community alone accounting for substantial (approximately 20%) variation in total carbon mineralized. Communities that shared a common history with a given foliar litter exhibited higher decomposition rates when compared to communities foreign to that habitat. Our results suggest that the implicit assumption in ecosystem models (i.e., microbial communities in the same environment are functionally equivalent) is incorrect. To predict accurately how biogeochemical processes will respond to global change may require consideration of the community composition and/or adaptation of microbial communities to past resource environments.
Lianas (woody vines) are an important and dynamic component of many forests throughout the world, and increases in CO2, mean winter temperature, and forest fragmentation may promote their growth and proliferation in temperate forests. In this study, we used a 45-year data set to test the hypothesis that lianas have increased in abundance and basal area in the interiors of 14 deciduous temperate forests in Wisconsin (USA) since 1959. We also censused woody plants along a gradient from the forest edge to the interior in seven of these forests to test the hypothesis that the abundance of lianas declines significantly with increasing distance from the forest edge. We found that lianas did not increase in abundance within the interiors of temperate forests in Wisconsin over the last 45 years. However, relative and absolute liana abundance decreased sharply with increasing distance from forest edges. Our findings suggest that forest fragmentation, not climate change, may be increasing the abundance of lianas in northern deciduous temperate forests, and that lianas may further increase in abundance if the severity of forest fragmentation intensifies.
Terrestrial biosphere-atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle's response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100-200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing >80% of their initial mass in <300 days. High organic matter solubility throughout decomposition combined with high rainfall support a model of litter-layer decomposition in these rain forests in which rapid mass loss in the litter layer is dominated by leaching of dissolved organic matter (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil.
In January 1958, a survey of alpine flora was conducted along a recently constructed access road across the upper volcanic slopes of Mauna Loa, Hawaii (2525-3397 m). Only five native Hawaiian species were encountered on sparsely vegetated historic and prehistoric lava flows adjacent to the roadway. A resurvey of roadside flora in 2008 yielded a more than fourfold increase to 22 species, including nine native species not previously recorded. Eight new alien species have now invaded this alpine environment, although exclusively limited to a few individuals in ruderal habitat along the roadway. Alternative explanations for species invasion and altitudinal change over the past 50 years are evaluated: (1) changes related to continuing primary succession on ameliorating (weathering) young lava substrates; (2) local climate change; and (3) road improvements and increased vehicular access which promote enhanced car-borne dispersal of alien species derived from the expanding pool of potential colonizers naturalized on the island in recent decades. Unlike alpine environments in temperate latitudes, the energy component (warming) in climate change on Mauna Loa does not appear to be the unequivocal driver of plant invasion and range extension. Warming may be offset by other climate change factors including rainfall and evapotranspiration.
Invertebrate herbivores can impact plant performance and plant communities. Conversely, plants can affect the ability of herbivores to find, choose, and consume them through their functional traits. While single plant traits have been related to rates of herbivory, most often involving single herbivore-plant pairs, much less is known about which suite of plant traits is important for determining herbivory for a pool of plant species interacting with a natural herbivore community. In this study we measured aboveground herbivore damage on 51 herbaceous species growing in monocultures of a grassland biodiversity experiment and collected 42 different plant traits representing four trait groups: physiological, morphological, phenological, and herbivore related. Using the method of random forests and multiple regression, we identified seven traits that are important predictors of herbivore damage (leaf nitrogen and lignin concentration, number of coleopteran and hemipteran herbivores potentially feeding on the plants, leaf life span, stem growth form, and root architecture); leaf nitrogen and lignin concentration were the two most important predictors. The final model accounted for 63% of the variation in herbivore damage. Traits from all four trait groups were selected, showing that a variety of plant characteristics can be statistically important when assessing folivory, including root traits. Our results emphasize that it is necessary to use a multivariate approach for identifying traits affecting complex ecological processes such as herbivory.
We compared the leaf traits and plant performance of 53 co-occurring tree species in a semi-evergreen tropical moist forest community. The species differed in all leaf traits analyzed: leaf life span varied 11-fold among species, specific leaf area 5-fold, mass-based nitrogen 3-fold, mass-based assimilation rate 13-fold, mass-based respiration rate 15-fold, stomatal conductance 8-fold, and photosynthetic water use efficiency 4-fold. Photosynthetic traits were strongly coordinated, and specific leaf area predicted mass-based rates of assimilation and respiration; leaf life span predicted many other leaf characteristics. Leaf traits were closely associated with growth, survival, and light requirement of the species. Leaf investment strategies varied on a continuum trading off short-term carbon gain against long-term leaf persistence that, in turn, is linked to variation in whole-plant growth and survival. Leaf traits were good predictors of plant performance, both in gaps and in the forest understory. High growth in gaps is promoted by cheap, short-lived, and physiologically active leaves. High survival in the forest understory is enhanced by the formation of long-lived well protected leaves that reduce biomass loss by herbivory, mechanical disturbance, or leaf turnover. Leaf traits underlay this growth-survival trade-off; species with short-lived, physiologically active leaves have high growth but low survival. This continuum in leaf traits, through its effect on plant performance, in turn gives rise to a continuum in species' light requirements.
Tree architecture is an important determinant of the height extension, light capture, and mechanical stability of trees, and it allows species to exploit the vertical height gradient in the forest canopy and horizontal light gradients at the forest floor. Tropical tree species partition these gradients through variation in adult stature (Hmax) and light demand. In this study we compare 22 architectural traits for 54 Bolivian moist-forest tree species. We evaluate how architectural traits related to Hmax vary with tree size, and we present a conceptual scheme in which we combine the two axes into four different functional groups. Interspecific correlations between architecture and Hmax varied strongly from negative to positive, depending on the reference sizes used. Stem height was positively related to Hmax at larger reference diameters (14-80 cm). Species height vs. diameter curves often flattened toward their upper ends in association with reproductive maturity for species of all sizes. Thus, adult understory trees were typically shorter than similar-diameter juveniles of larger species. Crown area was negatively correlated with Hmax at small reference heights and positively correlated at larger reference heights (15-34 m). Wide crowns allow the small understory species to intercept light over a large area at the expense of a reduced height growth. Crown length was negatively correlated with Hmax at intermediate reference heights (4-14 m). A long crown enables small understory species to maximize light interception in a light-limited environment. Light-demanding species were characterized by orthotropic stems and branches, large leaves, and a monolayer leaf arrangement. They realized an efficient height growth through the formation of narrow and shallow crowns. Light demand turned out to be a much stronger predictor of tree architecture than Hmax, probably because of the relatively low, open, and semi-evergreen canopy at the research site. The existence of four functional groups (shade-tolerant, partial-shade-tolerant, and long- and short-lived pioneer) was confirmed by the principal component and discriminant analysis. Both light demand and Hmax capture the major variation in functional traits found among tropical rain forest tree species, and the two-way classification scheme provides a straightforward model to understand niche differentiation in tropical forests.
Fungal endophytes are found in asymptomatic photosynthetic tissues of all major lineages of land plants. The ubiquity of these cryptic symbionts is clear, but the scale of their diversity, host range, and geographic distributions are unknown. To explore the putative hyperdiversity of tropical leaf endophytes, we compared endophyte communities along a broad latitudinal gradient from the Canadian arctic to the lowland tropical forest of central Panama. Here, we use molecular sequence data from 1403 endophyte strains to show that endophytes increase in incidence, diversity, and host breadth from arctic to tropical sites. Endophyte communities from higher latitudes are characterized by relatively few species from many different classes of Ascomycota, whereas tropical endophyte assemblages are dominated by a small number of classes with a very large number of endophytic species. The most easily cultivated endophytes from tropical plants have wide host ranges, but communities are dominated by a large number of rare species whose host range is unclear. Even when only the most easily cultured species are considered, leaves of tropical trees represent hotspots of fungal species diversity, containing numerous species not yet recovered from other biomes. The challenge remains to recover and identify those elusive and rarely cultured taxa with narrower host ranges, and to elucidate the ecological roles of these little-known symbionts in tropical forests.
Ecological processes of low-productivity ecosystems have long been considered to be driven by abiotic controls with biotic interactions playing an insignificant role. However, existing studies present conflicting evidence concerning the roles of these factors, in part due to the short temporal extent of most data sets and inability to test indirect effects of environmental variables modulated by biotic interactions. Using structural equation modeling to analyze 65 years of perennial vegetation change in the Sonoran Desert, we found that precipitation had a stronger positive effect on recruitment beneath existing canopies than in open microsites due to reduced evaporation rates. Variation in perennial canopy cover had additional facilitative effects on juvenile recruitment, which was indirectly driven by effects of density and precipitation on cover. Mortality was strongly influenced by competition as indicated by negative density-dependence, whereas precipitation had no effect. The combined direct, indirect, and interactive facilitative effects of precipitation and cover on recruitment were substantial, as was the effect of competition on mortality, providing strong evidence for dual control of community dynamics by climate and biotic interactions. Through an empirically derived simulation model, we also found that the positive feedback of density on cover produces unique temporal abundance patterns, buffering changes in abundance from high frequency variation in precipitation, amplifying effects of low frequency variation, and decoupling community abundance from precipitation patterns at high abundance. Such dynamics should be generally applicable to low-productivity systems in which facilitation is important and can only be understood within the context of long-term variation in climatic patterns. This predictive model can be applied to better manage low-productivity ecosystems, in which variation in biogeochemical processes and trophic dynamics may be driven by positive density-dependent feedbacks that influence temporal abundance and productivity patterns.
Recent observations and model simulations have highlighted the sensitivity of the forest-tundra ecotone to climatic forcing. In contrast, paleoecological studies have not provided evidence of tree-line fluctuations in response to Holocene climatic changes in Alaska, suggesting that the forest-tundra boundary in certain areas may be relatively stable at multicentennial to millennial time scales. We conducted a multiproxy study of sediment cores from an Alaskan lake near the altitudinal limits of key boreal-forest species. Paleoecological data were compared with independent climatic reconstructions to assess ecosystem responses of the forest tundra boundary to Little Ice Age (LIA) climatic fluctuations. Pollen, diatom, charcoal, macrofossil, and magnetic analyses provide the first continuous record of vegetation fire-climate interactions at decadal to centennial time scales during the past 700 years from southern Alaska. Boreal-forest diebacks characterized by declines of Picea mariana, P. glauca, and tree Betula occurred during the LIA (AD 1500-1800), whereas shrubs (Alnus viridis, Betula glandulosa/nana) and herbaceous taxa (Epilobium, Aconitum) expanded. Marked increases in charcoal abundance and changes in magnetic properties suggest increases in fire importance and soil erosion during the same period. In addition, the conspicuous reduction or disappearance of certain aquatic (e.g., Isoetes, Nuphar, Pediastrum) and wetland (Sphagnum) plants and major shifts in diatom assemblages suggest pronounced lake-level fluctuations and rapid ecosystem reorganization in response to LIA climatic deterioration. Our results imply that temperature shifts of 1-2 degrees C, when accompanied by major changes in moisture balance, can greatly alter high-altitudinal terrestrial, wetland, and aquatic ecosystems, including conversion between boreal-forest tree line and tundra. The climatic and ecosystem variations in our study area appear to be coherent with changes in solar irradiance, suggesting that changes in solar activity contributed to the environmental instability of the past 700 years.