Invader removal triggers competitive release in a threatened avian predator.

Changes in the distribution and abundance of invasive species can have far-reaching ecological consequences. Programs to control invaders are common but gauging the effectiveness of such programs using carefully controlled, large-scale field experiments is rare, especially at higher trophic levels. Experimental manipulations coupled with long-term demographic monitoring can reveal the mechanistic underpinnings of interspecific competition among apex predators and suggest mitigation options for invasive species. We used a large-scale before-after control-impact removal experiment to investigate the effects of an invasive competitor, the barred owl (Strix varia), on the population dynamics of an iconic old-forest native species, the northern spotted owl (Strix occidentalis caurina). Removal of barred owls had a strong, positive effect on survival of sympatric spotted owls and a weaker but positive effect on spotted owl dispersal and recruitment. After removals, the estimated mean annual rate of population change for spotted owls stabilized in areas with removals (0.2% decline per year), but continued to decline sharply in areas without removals (12.1% decline per year). The results demonstrated that the most substantial changes in population dynamics of northern spotted owls over the past two decades were associated with the invasion, population expansion, and subsequent removal of barred owls. Our study provides experimental evidence of the demographic consequences of competitive release, where a threatened avian predator was freed from restrictions imposed on its population dynamics with the removal of a competitively dominant invasive species.

Controlling biases in targeted plant removal experiments.

Targeted removal experiments are a powerful tool to assess the effects of plant species or (functional) groups on ecosystem functions. However, removing plant biomass in itself can bias the observed responses. This bias is commonly addressed by waiting until ecosystem recovery, but this is inherently based on unverified proxies or anecdotal evidence. Statistical control methods are efficient, but restricted in scope by underlying assumptions. We propose accounting for such biases within the experimental design, using a gradient of biomass removal controls. We demonstrate the relevance of this design by presenting (1) conceptual examples of suspected biases and (2) how to observe and control for these biases. Using data from a mycorrhizal association-based removal experiment, we show that ignoring biomass removal biases (including by assuming ecosystem recovery) can lead to incorrect, or even contrary conclusions (e.g. false positive and false negative). Our gradient design can prevent such incorrect interpretations, regardless of whether aboveground biomass has fully recovered. Our approach provides more objective and quantitative insights, independently assessed for each variable, than using a proxy to assume ecosystem recovery. Our approach circumvents the strict statistical assumptions of, for example, ANCOVA and thus offers greater flexibility in data analysis.

Consequences of the Reproductive Effort of Dioecious Taxus baccata L. Females in a Generative Bud Removal Experiment-Important Role of Nitrogen in Female Reproduction.

Dioecious species differ in the pattern and intensity of male and female reproductive investments. We aimed to determine whether female shoots deprived of generative buds show biochemical features, indicating their less-pronounced reproductive effort. For this purpose, the same branches of mature Taxus baccata females were deprived of generative organs. In the second and third years of the experiment, measurements were made in every season from the control and bud-removed shoots of females and control males. Bud removal caused an increase in nitrogen concentration almost to the level detected in the needles of male specimens, but only in current-year needles. Moreover, differences between male and control female shoots were present in the C:N ratio and increment biomass, but they disappeared when bud removal was applied to females. Additionally, between-sex differences were observed for content of phenolic compounds, carbon and starch, and SLA, independent of the female shoot reproductive effort. The study revealed that nitrogen uptake in seeds and arils may explain the lower nitrogen level and consequently the lower growth rate of females compared to males. At the same time, reproduction did not disturb carbon level in adjacent tissues, and two hypotheses explaining this phenomenon have been put forward.

Lianas do not reduce tree biomass accumulation in young successional tropical dry forests.

Young successional tropical forests are crucial in the global carbon cycle because they can quickly sequester large quantities of atmospheric carbon. However, lianas (woody vines) can significantly decrease biomass accumulation in young regenerating forests. Lianas are abundant in tropical dry forests, and thus we hypothesized that lianas reduce biomass accretion in dry forests. Lianas may be particularly detrimental to the growth of young trees, which are vulnerable to competition from lianas. Alternatively, lianas may have a stronger negative effect on the largest trees because lianas seek the high-light environment at the top of the forest canopy. We tested these hypotheses using a liana-removal experiment in 13 dry forest stands that ranged from 1 to 70 years in southwestern Panama. We measured biomass accumulation annually for more than 10,000 stems from 2013 to 2017. Contrary to our expectations, liana removal had no effect on tree biomass accumulation across our successional forests and throughout our study period. Liana removal did not benefit smaller trees or larger trees. Lianas did not increase biomass accumulation on recruits, and did not increase biomass loss due to mortality. Surprisingly, removing lianas had a negative effect on three out of 41 tree species. Lianas had no effect on biomass accumulation and loss, possibly because: (1) trees allocated resources to roots instead of stems, (2) trees and lianas partitioned water, (3) higher irradiance after liana removal reduced soil moisture, or (4) low water availability might have been such a strong stressor that it reduced plant-plant competition.

Weak interspecific interactions in a sagebrush steppe? Conflicting evidence from observations and experiments.

Stable coexistence requires intraspecific limitations to be stronger than interspecific limitations. The greater the difference between intra- and interspecific limitations, the more stable the coexistence, and the weaker the competitive release any species should experience following removal of competitors. We conducted a removal experiment to test whether a previously estimated model, showing surprisingly weak interspecific competition for four dominant species in a sagebrush steppe, accurately predicts competitive release. Our treatments were (1) removal of all perennial grasses and (2) removal of the dominant shrub, Artemisia tripartita. We regressed survival, growth, and recruitment on the locations, sizes, and species identities of neighboring plants, along with an indicator variable for removal treatment. If our "baseline" regression model, which accounts for local plant-plant interactions, accurately explains the observed responses to removals, then the removal coefficient should be non-significant. For survival, the removal coefficients were never significantly different from zero, and only A. tripartita showed a (negative) response to removals at the recruitment stage. For growth, the removal treatment effect was significant and positive for two species, Poa secunda and Pseudoroegneria spicata, indicating that the baseline model underestimated interspecific competition. For all three grass species, population models based on the vital rate regressions that included removal effects projected 1.4- to 3-fold increases in equilibrium population size relative to the baseline model (no removal effects). However, we found no evidence of higher response to removal in quadrats with higher pretreatment cover of A. tripartita, or by plants experiencing higher pre-treatment crowding by A. tripartita, raising questions about the mechanisms driving the positive response to removal. While our results show the value of combining observations with a simple removal experiment, more tightly controlled experiments focused on underlying mechanisms may be required to conclusively validate or reject predictions from phenomenological models.

Stress in biological invasions: Introduced invasive grey squirrels increase physiological stress in native Eurasian red squirrels.

Invasive alien species can cause extinction of native species through processes including predation, interspecific competition for resources or disease-mediated competition. Increases in stress hormones in vertebrates may be associated with these processes and contribute to the decline in survival or reproduction of the native species. Eurasian red squirrels (Sciurus vulgaris) have gone extinct across much of the British Isles and parts of Northern Italy following the introduction of North American invasive grey squirrels (Sciurus carolinensis). We extracted glucocorticoid metabolites from faecal samples to measure whether the presence of the invasive species causes an increase in physiological stress in individuals of the native species. We show that native red squirrels in seven sites where they co-occurred with invasive grey squirrels had glucocorticoid concentrations that were three times higher than those in five sites without the invasive species. Moreover, in a longitudinal study, stress hormones in native red squirrels increased after colonisation by grey squirrels. When we experimentally reduced the abundance of the invasive grey squirrels, the concentration of faecal glucocorticoid metabolites in co-occurring red squirrels decreased significantly between pre- and postremoval periods. Hence, we found that the invasive species acts as a stressor which significantly increases the concentrations of glucocorticoids in the native species. Given that sustained elevations in glucocorticoids could reduce body growth and reproductive rate, our results are consistent with previous studies where the co-occurrence of the invasive grey squirrel was associated with smaller size and lower reproductive output in red squirrels.

Management of invasive populations of the freshwater crayfish Procambarus clarkii (Decapoda, Cambaridae): test of a population-control method and proposal of a standard monitoring approach.

Invasive species are one of the main threats to biodiversity. When an alien species is introduced into a new environment, fast identification and definition of management strategies may avoid or minimize impacts. When an invasive species is already established, the most adopted approaches are population control and monitoring. In order to perform such strategies, assessment of characteristics of the invasive population is imperative. This study tested a new method of population size estimation and monitoring in an invasive population of crayfish Procambarus clarkii in a conservation area in the Atlantic Rain Forest (Southeastern Brazil). The population dynamics was studied for 1 year to examine the efficacy of the selected method and to evaluate if the population is stable. Later, the effect of periodical removal of animals on the population size was tested. The method of population estimation used in this study proved to be very effective. We recommend using it to monitor invasive populations of P. clarkii. The population size varied discretely over the year with variable but low growth rate, indicating that the population is already established which introduce a notable threat to native species. The continuous removal of specimens proved to be inefficient since the growth rate was higher after the removal. One intensive removal event might be more effective than a continuous moderate removal as the one applied in this study.

Effects of functional diversity loss on ecosystem functions are influenced by compensation.

Understanding the impacts of biodiversity loss on ecosystem functioning and services has been a central issue in ecology. Experiments in synthetic communities suggest that biodiversity loss may erode a set of ecosystem functions, but studies in natural communities indicate that the effects of biodiversity loss are usually weak and that multiple functions can be sustained by relatively few species. Yet, the mechanisms by which natural ecosystems are able to maintain multiple functions in the face of diversity loss remain poorly understood. With a long-term and large-scale removal experiment in the Inner Mongolian grassland, here we showed that losses of plant functional groups (PFGs) can reduce multiple ecosystem functions, including biomass production, soil NO3 -N use, net ecosystem carbon exchange, gross ecosystem productivity, and ecosystem respiration, but the magnitudes of these effects depended largely on which PFGs were removed. Removing the two dominant PFGs (perennial rhizomatous grasses and perennial bunchgrasses) simultaneously resulted in dramatic declines in all examined functions, but such declines were circumvented when either dominant PFG was present. We identify the major mechanism for this as a compensation effect by which each dominant PFG can mitigate the losses of others. This study provides evidence that compensation ensuing from PFG losses can mitigate their negative consequence, and thus natural communities may be more resilient to biodiversity loss than currently thought if the remaining PFGs have strong compensation capabilities. On the other hand, ecosystems without well-developed compensatory functional diversity may be much more vulnerable to biodiversity loss.

From facilitation to competition: temperature-driven shift in dominant plant interactions affects population dynamics in seminatural grasslands.

Biotic interactions are often ignored in assessments of climate change impacts. However, climate-related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co-occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad-scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population-level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.

Plasticity in functional traits in the context of climate change: a case study of the subalpine forb Boechera stricta (Brassicaceae).

Environmental variation often induces shifts in functional traits, yet we know little about whether plasticity will reduce extinction risks under climate change. As climate change proceeds, phenotypic plasticity could enable species with limited dispersal capacity to persist in situ, and migrating populations of other species to establish in new sites at higher elevations or latitudes. Alternatively, climate change could induce maladaptive plasticity, reducing fitness, and potentially stalling adaptation and migration. Here, we quantified plasticity in life history, foliar morphology, and ecophysiology in Boechera stricta (Brassicaceae), a perennial forb native to the Rocky Mountains. In this region, warming winters are reducing snowpack and warming springs are advancing the timing of snow melt. We hypothesized that traits that were historically advantageous in hot and dry, low-elevation locations will be favored at higher elevation sites due to climate change. To test this hypothesis, we quantified trait variation in natural populations across an elevational gradient. We then estimated plasticity and genetic variation in common gardens at two elevations. Finally, we tested whether climatic manipulations induce plasticity, with the prediction that plants exposed to early snow removal would resemble individuals from lower elevation populations. In natural populations, foliar morphology and ecophysiology varied with elevation in the predicted directions. In the common gardens, trait plasticity was generally concordant with phenotypic clines from the natural populations. Experimental snow removal advanced flowering phenology by 7 days, which is similar in magnitude to flowering time shifts over 2-3 decades of climate change. Therefore, snow manipulations in this system can be used to predict eco-evolutionary responses to global change. Snow removal also altered foliar morphology, but in unexpected ways. Extensive plasticity could buffer against immediate fitness declines due to changing climates.

Competitive exclusion, beta diversity, and deterministic vs. stochastic drivers of community assembly.

Species diversity has two components - number of species and spatial turnover in species composition (beta-diversity). Using a field experiment focusing on a system of Mediterranean grasslands, we show that interspecific competition may influence the two components in the same direction or in opposite directions, depending on whether competitive exclusions are deterministic or stochastic. Deterministic exclusions reduce both patch-scale richness and beta-diversity, thereby homogenising the community. Stochastic extinctions reduce richness at the patch scale, but increase the differences in species composition among patches. These results indicate that studies of competitive effects on beta diversity may help to distinguish between deterministic and stochastic components of competitive exclusion. Such distinction is crucial for understanding the causal relationship between competition and species diversity, one of the oldest and most fundamental questions in ecology.

Experimental induction of social instability during early breeding does not alter testosterone levels in male black redstarts, a socially monogamous songbird.

Testosterone plays an important role in territorial behavior of many male vertebrates and the Challenge Hypothesis has been suggested to explain differences in testosterone concentrations between males. For socially monogamous birds, the challenge hypothesis predicts that testosterone should increase during male-male interactions. To test this, simulated territorial intrusion (STI) experiments have been conducted, but only about a third of all bird species investigated so far show the expected increase in testosterone. Previous studies have shown that male black redstarts (Phoenicurus ochruros) do not increase testosterone during STIs or short-term male-male challenges. The aim of this study was to evaluate whether black redstarts modulate testosterone in an experimentally induced longer-term unstable social situation. We created social instability by removing males from their territories and compared the behavior and testosterone concentrations of replacement males and neighbors with those of control areas. Testosterone levels did not differ among replacement males, neighbors and control males. Injections with GnRH resulted in elevation of testosterone in all groups, suggesting that all males were capable of increasing testosterone. We found no difference in the behavioral response to STIs between control and replacement males. Furthermore, there was no difference in testosterone levels between replacement males that had expanded their territory and new-coming males. In combination with prior work these data suggest that testosterone is not modulated by male-male interactions in black redstarts and that testosterone plays only a minor role in territorial behavior. We suggest that territorial behavior in species that are territorial throughout most of their annual life-cycle may be decoupled from testosterone.

Supra-optimal leaf area index of a temperate liana Pueraria lobata for competition with Solidago altissima at the expense of canopy photosynthesis.

Leaf area index (LAI) measured for the actual plant canopy is higher than the LAI that maximizes canopy photosynthesis (referred to as optimal LAI) because each individual can increase its light interception by unilaterally producing more leaf area. The LAI of an invasive woody vine Pueraria lobata (kudzu) is one of the highest among plant species, sometimes attaining nearly 10 m2 m-2. The high LAI casts heavy shade over neighboring plants, making their survival difficult. Interesting to note is that the high LAI also increases self-shading, thereby decreasing its own photosynthesis processes. In the present study, the influences of the high LAI on light interception and canopy photosynthesis, as well as on the inter-specific competition was investigated on a roadside P. lobata vegetation in Japan. With the aid of a canopy photosynthesis model and a sensitivity analysis, it was revealed that the actual LAI was 2.2-3.0 times higher than the optimal LAI for maximizing canopy photosynthesis. In the following year, a field experiment was conducted where a nearly optimal LAI was maintained throughout the growth period by regularly clipping the leaves of P. lobata. Ultimately, the field results revealed that even with a nearly optimal LAI, P. lobata was outcompeted by a competing alien weed, Solidago altissima (tall goldenrod). These results indicate that the supra-optimal leaf area, rather than maximum canopy carbon gain, makes P. lobata the dominating species in light-competing environments.

Invasive Impatiens glandulifera: A driver of changes in native vegetation?

Biological invasions are one of the major threats to biodiversity worldwide and contribute to changing community patterns and ecosystem processes. However, it is often not obvious whether an invader is the "driver" causing ecosystem changes or a "passenger" which is facilitated by previous ecosystem changes. Causality of the impact can be demonstrated by experimental removal of the invader or introduction into a native community. Using such an experimental approach, we tested whether the impact of the invasive plant Impatiens glandulifera on native vegetation is causal, and whether the impact is habitat-dependent. We conducted a field study comparing invaded and uninvaded plots with plots from which I. glandulifera was removed and plots where I. glandulifera was planted within two riparian habitats, alder forests and meadows. A negative impact of planting I. glandulifera and a concurrent positive effect of removal on the native vegetation indicated a causal effect of I. glandulifera on total native biomass and growth of Urtica dioica. Species α-diversity and composition were not affected by I. glandulifera manipulations. Thus, I. glandulifera had a causal but low effect on the native vegetation. The impact depended slightly on habitat as only the effect of I. glandulifera planting on total biomass was slightly stronger in alder forests than meadows. We suggest that I. glandulifera is a "back-seat driver" of changes, which is facilitated by previous ecosystem changes but is also a driver of further changes. Small restrictions of growth of the planted I. glandulifera and general association of I. glandulifera with disturbances indicate characteristics of a back-seat driver. For management of I. glandulifera populations, this requires habitat restoration along with removal of the invader.

Withering corolla remains on the calyx tube enhance reproductive success in Oxalis stricta L.

Several flower traits can affect plant reproductive fitness via pollinator attraction, herbivory defense, and thermal regulation of the pistil. In this study, we focus on thermal regulation of the pistil after flowering. We experimentally investigated the functional significance of the withering corollas that remain attached to the calyx tubes of Oxalis stricta L. We studied thermal regulation of the pistil by removing corollas and comparing the plants with and without corollas, under regulated dark and light periods, with an ambient temperature during the dark period lower than that during the light period. In plants lacking corollas, the pistil temperature was significantly lower than in control plants (with intact corollas) by approximately 2°C. Although fruit set in the corolla-removed plants was not significantly different from that in control plants, the temperature threshold for 50% fruition in the corolla-removed plants was significantly higher than that in the controls. Furthermore, the seed number, total seed weight, and single-grain weight were significantly lower in the corolla-removed plants than in control plants. The estimated annual number of reproductive cycles (from June to October), total seed number, and total seed weights were also lower in corolla-removed plants. These findings indicate that the withering corolla remains play a role in thermoregulation of the pistil, and thereby enhance reproductive success. Our study is the first to validate one of the assumed ecological roles of the withering remains of plant corollas.

Competition for safe real estate, not food, drives density-dependent juvenile survival in a large herbivore.

Density-dependent competition for food reduces vital rates, with juvenile survival often the first to decline. A clear prediction of food-based, density-dependent competition for large herbivores is decreasing juvenile survival with increasing density. However, competition for enemy-free space could also be a significant mechanism for density dependence in territorial species. How juvenile survival is predicted to change across density depends critically on the nature of predator-prey dynamics and spatial overlap among predator and prey, especially in multiple-predator systems. Here, we used a management experiment that reduced densities of a generalist predator, coyotes, and specialist predator, mountain lions, over a 5-year period to test for spatial density dependence mediated by predation on juvenile mule deer in Idaho, USA. We tested the spatial density-dependence hypothesis by tracking the fate of 251 juvenile mule deer, estimating cause-specific mortality, and testing responses to changes in deer density and predator abundance. Overall juvenile mortality did not increase with deer density, but generalist coyote-caused mortality did, but not when coyote density was reduced experimentally. Mountain lion-caused mortality did not change with deer density in the reference area in contradiction of the food-based competition hypothesis, but declined in the treatment area, opposite to the pattern of coyotes. These observations clearly reject the food-based density-dependence hypothesis for juvenile mule deer. Instead, our results provide support for the spatial density-dependence hypothesis that competition for enemy-free space increases predation by generalist predators on juvenile large herbivores.

Requirements for the spatial storage effect are weakly evident for common species in natural annual plant assemblages.

Coexistence in spatially varying environments is theorized to be promoted by a variety of mechanisms including the spatial storage effect. The spatial storage effect promotes coexistence when (1) species have unique vital rate responses to their spatial environment and, when abundant, (2) experience stronger competition in the environmental patches where they perform better. In a naturally occurring southwest Western Australian annual plant system, we conducted a neighbor removal experiment involving eleven focal species growing in high-abundance populations. Specifically, we measured species' fecundity across a variety of environmental gradients in both the presence and absence of neighbors. For the environmental variables that we measured, there was only limited evidence for species-specific responses to the environment, with a composite variable describing overstory cover and leaf litter cover being the best predictor of fecundity for a subset of focal species. In addition, although we found strong evidence for intraspecific competition, positive environment-competition covariance was only detected for one species. Thus, positive environment-competition covariance may not be as common as expected in populations of species growing at high abundance, at least when tested in natural assemblages. Our findings highlight the inherent limitations of using natural assemblages to study spatial coexistence mechanisms, and we urge empirical ecologists to take these limitations into account when designing future experiments.

Lianas reduce biomass accumulation in early successional tropical forests.

Early successional tropical forests could mitigate climate change via rapid accumulation of atmospheric carbon. However, liana (woody vine) abundance and biomass has been increasing in many tropical forests over the past decades, which may slow the speed at which secondary forests accumulate biomass. Lianas decrease biomass accumulation in tropical forests, and may have a particularly strong effect on young forests by stalling tree growth. As forests mature, trees may outgrow or shed lianas, thus escaping some of the negative effects of lianas. Alternatively, lianas may have the strongest effect in older successional forests if the effect of lianas is commensurate with their density, which increases dramatically in the first decades of forest succession. We tested these two hypotheses using a landscape liana-removal experiment in 30 forest stands that ranged from 10 to 35 yr old in Central Panama. We measured tree growth and biomass accumulation in the stands every year from 2014 to 2017. We found that the effect of liana removal on large trees (≥20-cm diameter) decreased with forest age, supporting the hypothesis that lianas have the strongest negative effects on trees, and thus biomass uptake and carbon storage, in very young successional forests. Large trees accumulated more biomass in the absence of lianas in younger forests than in older forests (compared to controls) even after accounting for the effect of canopy completeness and crown illumination, implying that the detrimental effects of lianas go well beyond resource availability and crown health. There was no significant effect of lianas on small trees (1-20-cm diameter), likely because lianas seek light and thus do not deploy their leaves on small trees that are trapped in the forest understory. Our results show that high liana density early in forest succession reduces forest biomass accumulation by negatively impacting large trees, thus decreasing the capacity of young secondary forests to mitigate climate change. Although the negative effects of lianas on forest biomass diminish as forests age, they do not disappear, and thus lianas are an important component of tropical forest carbon budgets throughout succession.

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Plant-plant interactions play a key role in regulating the composition and structure of communities and ecosystems. Studies of plant-plant interactions in forest ecosystems have traditionally concentrated on either tree-tree interactions or overstory species' impacts on understory plants. The possible effects of understory species on overstory trees have received less attention. We summarized the effects of understory species on soil physiological properties, soil fauna activities, leaf litter decomposition, and ecophysiology and growth of the overstory species. Then the effects of distur-bance on understory-overstory interactions were discussed. Finally, an ecophysiology-based concept model of understory effects on overstory trees was proposed. Understory removal experiments showed that the study area, overstory species age, soil fertility and understory species could significantly affect the understory-overstory interactions.